text:
Chordate

A chordate is a deuterostomic animal belonging to the phylum Chordata. All chordates possess, at some point during their larval or adult stages, five distinctive physical characteristics (synapomorphies) that distinguish them from other taxa. These five synapomorphies are a notochord, a hollow dorsal nerve cord, an endostyle or thyroid, pharyngeal slits, and a post-anal tail. The name "chordate" comes from the first of these synapomorphies, the notochord, which plays a significant role in chordate body plan structuring and movements. Chordates are also bilaterally symmetric, have a coelom, possess a closed circulatory system, and exhibit metameric segmentation.

In addition to the morphological characteristics used to define chordates, analysis of genome sequences has identified two conserved signature indels (CSIs) in their proteins: cyclophilin-like protein and inner mitochondrial membrane protease ATP23, which are exclusively shared by all vertebrates, tunicates and cephalochordates. These CSIs provide molecular means to reliably distinguish chordates from all other animals.

Chordates are divided into three subphyla: Vertebrata (fish, amphibians, reptiles, birds and mammals), whose notochords are replaced by a cartilaginous/bony axial endoskeleton (spine) and are cladistically and phylogenetically a subgroup of the clade Craniata (i.e. chordates with a skull); Tunicata or Urochordata (sea squirts, salps, and larvaceans), which only retain the synapomorphies during their larval stage; and Cephalochordata (lancelets), which resemble fish but have no gills. The Vertebrates and Tunicates compose the clade Olfactores, which is sister to Cephalochordata (see diagram under Phylogeny). Extinct taxa such as the Conodonts are Chordata, but their internal placement is less certain. Hemichordata (which includes the acorn worms) was previously considered a fourth chordate subphylum, but now is treated as a separate phylum. Hemichordates are now thought to be closer to the Echinoderms, and together they form the clade Ambulacraria, the sister phylum of the Chordates. Chordata, Ambulacraria, and possibly Xenacoelomorpha are believed to form the superphylum Deuterostomia, although this has recently been called into doubt.

Chordates are the third-largest phylum of the animal kingdom (behind arthropods and molluscs) and is also one of the most ancient taxons. Chordate fossils have been found from as early as the Cambrian explosion over 539 million years ago. Of the more than 81,000 living species of chordates, about half are ray-finned fishes (class Actinopterygii) and the vast majority of the rest are tetrapods, a terrestrial clade of lobe-finned fishes (Sarcopterygii) who evolved air-breathing using lungs.

History of name
Although the name Chordata is attributed to William Bateson (1885), it was already in prevalent use by 1880. Ernst Haeckel described a taxon comprising tunicates, cephalochordates, and vertebrates in 1866. Though he used the German vernacular form, it is allowed under the ICZN code because of its subsequent latinization.

Anatomy
Chordates form a phylum of animals that are defined by having at some stage in their lives all of the following anatomical features:

 * A notochord, a stiff but elastic rod of glycoprotein wrapped in two collagen helices, which extends along the central axis of the body. Among members of the subphylum Vertebrata (vertebrates), the notochord gets replaced by hyaline cartilage or osseous tissue of the spine, and notochord remnants develop into the intervertebral discs, which allow adjacent spinal vertebrae to bend and twist relative to each other. In wholly aquatic species, this helps the animal swim efficiently by flexing its tail side-to-side.
 * A hollow dorsal nerve cord, also known as the neural tube, which develops into the spinal cord, the main communications trunk of the nervous system. In vertebrates, the rostral end of the neural tube enlarges into several vesicles during embryonic development, which give rise to the brain.
 * Pharyngeal slits. The pharynx is the part of the throat immediately behind the mouth. In fish, the slits are modified to form gills, but in some other chordates they are part of a filter-feeding system that extracts food particles from ingested water. In tetrapods, they are only present during embryonic stages of the development.
 * A post-anal tail. A muscular tail that extends backwards behind the anus. In some chordates such as hominids, this is only present in the embryonic stage.
 * An endostyle. This is a groove in the ventral wall of the pharynx. In filter-feeding species it produces mucus to gather food particles, which helps in transporting food to the esophagus. It also stores iodine, and may be a precursor of the vertebrate thyroid gland.

There are soft constraints that separate chordates from other biological lineages, but are not part of the formal definition:
 * All chordates are deuterostomes. This means that, during embryonic development, the anus forms before the mouth does.
 * All chordates are based on a bilateral body plan.
 * All chordates are coelomates, and have a fluid-filled body cavity (coelom) with a complete serosal lining derived from mesoderm called mesothelium (see Brusca and Brusca).

Classification
The following schema is from the 2015 edition of Vertebrate Palaeontology. The invertebrate chordate classes are from Fishes of the World. While it is structured so as to reflect evolutionary relationships (similar to a cladogram), it also retains the traditional ranks used in Linnaean taxonomy.

 * Phylum Chordata
 * Subphylum Cephalochordata (Acraniata) – (lancelets; 30 species)
 * Class Leptocardii (lancelets)
 * Clade Olfactores
 * Subphylum Tunicata (Urochordata) – (tunicates; 3,000 species)
 * Class Ascidiacea (sea squirts)
 * Class Thaliacea (salps, doliolids and pyrosomes)
 * Class Appendicularia (larvaceans)
 * Class Sorberacea
 * Subphylum Vertebrata (Craniata) (vertebrates – animals with backbones; 66,100+ species)
 * Superclass 'Agnatha' paraphyletic (jawless vertebrates; 100+ species)
 * Class Cyclostomata
 * Infraclass Myxinoidea or Myxini (hagfish; 65 species)
 * Infraclass Petromyzontida or Hyperoartia (lampreys)
 * Class †Conodonta
 * Class †Myllokunmingiida
 * Class †Pteraspidomorphi
 * Class †Thelodonti
 * Class †Anaspida
 * Class †Cephalaspidomorphi
 * Infraphylum Gnathostomata (jawed vertebrates)
 * Class †Placodermi (Paleozoic armoured forms; paraphyletic in relation to all other gnathostomes)
 * Class Chondrichthyes (cartilaginous fish; 900+ species)
 * Class †Acanthodii (Paleozoic "spiny sharks"; paraphyletic in relation to Chondrichthyes)
 * Class Osteichthyes (bony fish; 30,000+ species)
 * Subclass Actinopterygii (ray-finned fish; about 30,000 species)
 * Subclass Sarcopterygii (lobe-finned fish: 8 species)
 * Superclass Tetrapoda (four-limbed vertebrates; 35,100+ species) (The classification below follows Benton 2004, and uses a synthesis of rank-based Linnaean taxonomy and also reflects evolutionary relationships. Benton included the Superclass Tetrapoda in the Subclass Sarcopterygii in order to reflect the direct descent of tetrapods from lobe-finned fish, despite the former being assigned a higher taxonomic rank.)
 * Class Amphibia (amphibians; 8,100+ species)
 * Class Sauropsida (reptiles (including birds); 21,300+ species – 10,000+ species of birds and 11,300+ species of reptiles)
 * Class Synapsida (mammals; 5,700+ species)

Cephalochordata: Lancelets
Cephalochordates, one of the three subdivisions of chordates, are small, "vaguely fish-shaped" animals that lack brains, clearly defined heads and specialized sense organs. These burrowing filter-feeders compose the earliest-branching chordate subphylum.

Tunicata (Urochordata)

Most tunicates appear as adults in one of two major forms, known as "sea squirts" and salps. Both of these are soft-bodied filter-feeders that lack the standard features of chordates, which are only retained in their larvae. Sea squirts are sessile and consist mainly of water pumps and filter-feeding apparatus; salps float in mid-water, feeding on plankton, and have a two-generation cycle in which one generation is solitary and the next forms chain-like colonies. However, all tunicate larvae have the standard chordate features, including long, tadpole-like tails; they also have rudimentary brains, light sensors and tilt sensors. The third main group of tunicates, Appendicularia (also known as Larvacea), retain tadpole-like shapes and active swimming all their lives, and were for a long time regarded as larvae of sea squirts or salps. The etymology of the term Urochordata (Balfour 1881) is from the ancient Greek οὐρά (oura, "tail") + Latin chorda ("cord"), because the notochord is only found in the tail. The term Tunicata (Lamarck 1816) is recognised as having precedence and is now more commonly used.

Craniata (Vertebrata)

Craniates all have distinct skulls. They include the hagfish, which have no vertebrae. Michael J. Benton commented that "craniates are characterized by their heads, just as chordates, or possibly all deuterostomes, are by their tails".

Most craniates are vertebrates, in which the notochord is replaced by the vertebral column. It consists of a series of bony or cartilaginous cylindrical vertebrae, generally with neural arches that protect the spinal cord, and with projections that link the vertebrae. Hagfishes have incomplete braincases and no vertebrae, and are therefore not regarded as vertebrates, but they are members of the craniates, the group within which vertebrates are thought to have evolved. However the cladistic exclusion of hagfish from the vertebrates is controversial, as they may instead be degenerate vertebrates who have secondarily lost their vertebral columns.

The position of lampreys is ambiguous. They have complete braincases and rudimentary vertebrae, and therefore may be regarded as vertebrates and true fish. However, molecular phylogenetics, which uses biochemical features to classify organisms, has produced both results that group them with vertebrates and others that group them with hagfish. If lampreys are more closely related to the hagfish than the other vertebrates, this would suggest that they form a clade, which has been named the Cyclostomata.

Overview

There is still much ongoing differential (DNA sequence based) comparison research that is trying to separate out the simplest forms of chordates. As some lineages of the 90% of species that lack a backbone or notochord might have lost these structures over time, this complicates the classification of chordates. Some chordate lineages may only be found by DNA analysis, when there is no physical trace of any chordate-like structures.

Attempts to work out the evolutionary relationships of the chordates have produced several hypotheses. The current consensus is that chordates are monophyletic, meaning that the Chordata include all and only the descendants of a single common ancestor, which is itself a chordate, and that craniates' nearest relatives are tunicates. Recent identification of two conserved signature indels (CSIs) in the proteins cyclophilin-like protein and mitochondrial inner membrane protease ATP23, which are exclusively shared by all vertebrates, tunicates and cephalochordates also provide strong evidence of the monophyly of Chordata.

All of the earliest chordate fossils have been found in the Early Cambrian Chengjiang fauna, and include two species that are regarded as fish, which implies that they are vertebrates. Because the fossil record of early chordates is poor, only molecular phylogenetics offers a reasonable prospect of dating their emergence. However, the use of molecular phylogenetics for dating evolutionary transitions is controversial.

It has also proved difficult to produce a detailed classification within the living chordates. Attempts to produce evolutionary "family trees" shows that many of the traditional classes are paraphyletic.

Diagram of the evolutionary relationships of chordates

While this has been well known since the 19th century, an insistence on only monophyletic taxa has resulted in vertebrate classification being in a state of flux.

The majority of animals more complex than jellyfish and other Cnidarians are split into two groups, the protostomes and deuterostomes, the latter of which contains chordates. It seems very likely the Kimberella was a member of the protostomes. If so, this means the protostome and deuterostome lineages must have split some time before Kimberella appeared—at least, and hence well before the start of the Cambrian. The Ediacaran fossil Ernietta, from about, may represent a deuterostome animal.

Fossils of one major deuterostome group, the echinoderms (whose modern members include starfish, sea urchins and crinoids), are quite common from the start of the Cambrian,. The Mid Cambrian fossil Rhabdotubus johanssoni has been interpreted as a pterobranch hemichordate. Opinions differ about whether the Chengjiang fauna fossil Yunnanozoon, from the earlier Cambrian, was a hemichordate or chordate. Another fossil, Haikouella lanceolata, also from the Chengjiang fauna, is interpreted as a chordate and possibly a craniate, as it shows signs of a heart, arteries, gill filaments, a tail, a neural chord with a brain at the front end, and possibly eyes—although it also had short tentacles round its mouth. Haikouichthys and Myllokunmingia, also from the Chengjiang fauna, are regarded as fish. Pikaia, discovered much earlier (1911) but from the Mid Cambrian Burgess Shale (505 Ma), is also regarded as a primitive chordate. On the other hand, fossils of early chordates are very rare, since invertebrate chordates have no bones or teeth, and only one has been reported for the rest of the Cambrian.

The evolutionary relationships between the chordate groups and between chordates as a whole and their closest deuterostome relatives have been debated since 1890. Studies based on anatomical, embryological, and paleontological data have produced different "family trees". Some closely linked chordates and hemichordates, but that idea is now rejected. Combining such analyses with data from a small set of ribosome RNA genes eliminated some older ideas, but opened up the possibility that tunicates (urochordates) are "basal deuterostomes", surviving members of the group from which echinoderms, hemichordates and chordates evolved. Some researchers believe that, within the chordates, craniates are most closely related to cephalochordates, but there are also reasons for regarding tunicates (urochordates) as craniates' closest relatives.

Since early chordates have left a poor fossil record, attempts have been made to calculate the key dates in their evolution by molecular phylogenetics techniques—by analyzing biochemical differences, mainly in RNA. One such study suggested that deuterostomes arose before and the earliest chordates around. However, molecular estimates of dates often disagree with each other and with the fossil record, and their assumption that the molecular clock runs at a known constant rate has been challenged.

Traditionally, Cephalochordata and Craniata were grouped into the proposed clade "Euchordata", which would have been the sister group to Tunicata/Urochordata. More recently, Cephalochordata has been thought of as a sister group to the "Olfactores", which includes the craniates and tunicates. The matter is not yet settled.

A specific relationship between Vertebrates and Tunicates is also strongly supported by two CSIs found in the proteins predicted exosome complex RRP44 and serine palmitoyltransferase, that are exclusively shared by species from these two subphyla but not Cephalochordates, indicating Vertebrates are more closely related to Tunicates than Cephalochordates.

Cladogram
Below is a phylogenetic tree of the phylum. Lines of the cladogram show probable evolutionary relationships between both extinct taxa, which are denoted with a dagger (†), and extant taxa. Relatives of vertebrates are invertebrates. The positions (relationships) of the lancelets, tunicates, and craniates/vertebrates are based on the following studies:

Closest nonchordate relatives
The closest relatives of the Chordates are believed to be the Hemichordates and Echinodermata, which together form the Ambulacraria. The Chordata and Ambulacraria together form the superphylum Deuterostomia.

Hemichordates
Hemichordates ("half chordates") have some features similar to those of chordates: branchial openings that open into the pharynx and look rather like gill slits; stomochords, similar in composition to notochords, but running in a circle round the "collar", which is ahead of the mouth; and a dorsal nerve cord—but also a smaller ventral nerve cord.

There are two living groups of hemichordates. The solitary enteropneusts, commonly known as "acorn worms", have long proboscises and worm-like bodies with up to 200 branchial slits, are up to 2.5 m long, and burrow though seafloor sediments. Pterobranchs are colonial animals, often less than 1 mm long individually, whose dwellings are interconnected. Each filter feeds by means of a pair of branched tentacles, and has a short, shield-shaped proboscis. The extinct graptolites, colonial animals whose fossils look like tiny hacksaw blades, lived in tubes similar to those of pterobranchs.

Echinoderms

Echinoderms differ from chordates and their other relatives in three conspicuous ways: they possess bilateral symmetry only as larvae – in adulthood they have radial symmetry, meaning that their body pattern is shaped like a wheel; they have tube feet; and their bodies are supported by dermal skeletons made of calcite, a material not used by chordates. Their hard, calcified shells keep their bodies well protected from the environment, and these skeletons enclose their bodies, but are also covered by thin skins. The feet are powered by another unique feature of echinoderms, a water vascular system of canals that also functions as a "lung" and surrounded by muscles that act as pumps. Crinoids are typically sessile and look rather like flowers (hence the common name "sea lilies"), and use their feather-like arms to filter food particles out of the water; most live anchored to rocks, but a few species can move very slowly. Other echinoderms are mobile and take a variety of body shapes, for example starfish and brittle stars, sea urchins and sea cucumbers.

title: Chordate


text:
Charlize Theron

Charlize Theron (born 7 August 1975) is a South African and American actress and producer. One of the world's highest-paid actresses, she is the recipient of various accolades, including an Academy Award and a Golden Globe Award. In 2016, Time named her one of the 100 most influential people in the world.

Theron came to international prominence in the 1990s by playing the leading lady in the Hollywood films The Devil's Advocate (1997), Mighty Joe Young (1998), and The Cider House Rules (1999). She received critical acclaim for her portrayal of serial killer Aileen Wuornos in Monster (2003), for which she won the Silver Bear and Academy Award for Best Actress, becoming the first South African to win an acting Oscar. She received another Academy Award nomination for playing a sexually abused woman seeking justice in the drama North Country (2005).

Theron has starred in several commercially successful action films, including The Italian Job (2003), Hancock (2008), Snow White and the Huntsman (2012), Prometheus (2012), Mad Max: Fury Road (2015), The Fate of the Furious (2017), Atomic Blonde (2017), The Old Guard (2020), F9 (2021) and Fast X (2023). She received praise for playing troubled women in Jason Reitman's comedy-dramas Young Adult (2011) and Tully (2018), and for portraying Megyn Kelly in the biographical drama Bombshell (2019), for which she received her third Academy Award nomination.

Since the early 2000s, Theron has ventured into film production with her company Denver and Delilah Productions. She has produced numerous films, in many of which she had a starring role, including The Burning Plain (2008), Dark Places (2015), and Long Shot (2019). Theron became an American citizen in 2007, while retaining her South African citizenship. She has been honoured with a motion picture star on the Hollywood Walk of Fame.

Early life
Theron was born in Benoni, in Transvaal Province (Gauteng Province since 1994) of South Africa on 7 August 1975. She is the only child of road constructionists Gerda (née Maritz) and Charles Theron (27 November 1947 – 21 June 1991). The Second Boer War military leader Danie Theron was her great-granduncle. She is from an Afrikaner family, and her ancestry includes Dutch as well as French and German. Her French forebears were early Huguenots in South Africa.

She grew up on her parents' farm in Benoni, near Johannesburg. On 21 June 1991, Theron's father, an alcoholic, threatened both Charlize and her mother while drunk, physically attacking her mother and firing a gun at both of them. Theron's mother retrieved her own handgun, shot back and killed him. The shooting was legally adjudged to have been self-defense, and her mother faced no charges.

Theron attended Putfontein Primary School (Laerskool Putfontein), a period during which she has said she was not "fitting in". She was frequently unwell with jaundice throughout childhood and the antibiotics she was administered made her upper incisor milk teeth rot; they had to be surgically removed. Theron's permanent teeth did not grow until she was roughly ten years old. At 13, Theron was sent to boarding school and began her studies at the National School of the Arts in Johannesburg. About her early life in her home country, Theron has said: “I grew up as an only child in South Africa, and there was turmoil in my family, but the surroundings were so great. I was usually barefoot in the dirt: no Game Boys, no computers, and we had sanctions, so there were no concerts. This meant you had to entertain yourself.”

Although Theron is fluent in English, her first language is Afrikaans.

Early work (1991–1996)
Although she saw herself as a dancer, Theron won a one-year modelling contract at age 16 at a local competition in Salerno, Italy and moved with her mother to Milan, Italy. After Theron spent a year modelling throughout Europe, she and her mother moved to the United States; they resided in New York City and Miami. In New York, she attended the Joffrey Ballet School, where she trained as a ballet dancer until a knee injury closed this career path. As Theron recalled in 2008:

"I went to New York for three days to model, and then I spent a winter in New York in a friend's windowless basement apartment. I was broke, I was taking class at the Joffrey Ballet, and my knees gave out. I realized I couldn't dance anymore, and I went into a major depression. My mom came over from South Africa and said, 'Either you figure out what to do next or you come home, because you can sulk in South Africa'."

In 1994, Theron flew to Los Angeles on a one-way ticket her mother bought for her; she intended to work in the film industry. During her initial months there, she lived in a motel with the $300 budget that her mother had given her; she continued receiving cheques from New York and lived "from paycheck to paycheck". Theron stole bread from a basket in a restaurant to survive. One day, she went to a Hollywood Boulevard bank to cash a few cheques, including one her mother had sent to help with the rent; however; the check from her mother was rejected because it was out-of-state and she was not an American citizen. Theron argued and pleaded with the bank teller until talent agent John Crosby, who was the next customer behind her, cashed it for her and gave her his business card.

Crosby introduced Theron to an acting school. In 1995, she played her first non-speaking film role in the horror film Children of the Corn III: Urban Harvest. In Theron's first speaking role, she portrayed hitwoman Helga Svelgen in 2 Days in the Valley (1996). Despite the film's mixed reviews, Theron drew attention due to her beauty and to a scene in which she fought Teri Hatcher's character. Theron feared being typecast as characters similar to Helga and recalled being asked to repeat her performance in the film during auditions: "A lot of people were saying, 'You should just hit while the iron's hot' [...] But playing the same part over and over doesn't leave you with any longevity. And I knew it was going to be harder for me, because of what I look like, to branch out to different kinds of roles".

When auditioning for Showgirls, Theron was introduced to talent agent J. J. Harris by co-casting director Johanna Ray. She recalled being surprised at how much faith Harris had in her potential. Theron has referred to Harris as her mentor. Harris found scripts and films for Theron in a variety of genres and encouraged her to become a producer. She served as Theron's agent for over 15 years.

Breakthrough (1997–2002)
Theron's career expanded by the end of the 1990s. In the horror drama The Devil's Advocate (1997), which is credited as her break-out film, Theron starred alongside Keanu Reeves and Al Pacino as the haunted wife of an unusually successful lawyer. She subsequently starred in the adventure film Mighty Joe Young (1998) as the friend and protector of a giant mountain gorilla, and in the drama The Cider House Rules (1999), as a woman who seeks an abortion in World War II-era Maine. While Mighty Joe Young flopped at the box office, The Devil's Advocate and The Cider House Rules were commercially successful. She appeared on the cover of the January 1999 issue of Vanity Fair as the "White Hot Venus". The May 1999 issue of Playboy magazine also featured Theron on its cover; the photos of Theron used by Playboy had been taken several years earlier when she was an unknown model, and Theron unsuccessfully sued the magazine for publishing them without her consent.

By the early 2000s, Theron continued to steadily take on roles in films such as Reindeer Games (2000), The Yards (2000), The Legend of Bagger Vance (2000), Men of Honor (2000), Sweet November (2001), The Curse of the Jade Scorpion (2001), and Trapped (2002), all of which, despite achieving only limited commercial success, helped to establish her as an actress. On this period in her career, Theron remarked: "I kept finding myself in a place where directors would back me but studios didn't. [I began] a love affair with directors, the ones I really, truly admired. I found myself making really bad movies, too. Reindeer Games was not a good movie, but I did it because I loved [director] John Frankenheimer".

Worldwide recognition and critical success (2003–2008)

Theron starred as a safe and vault technician in the 2003 heist film The Italian Job, an American homage/remake of the 1969 British film of the same name, directed by F. Gary Gray and opposite Mark Wahlberg, Edward Norton, Jason Statham, Seth Green, and Donald Sutherland. The film was a box office success, grossing US$176 million worldwide.

In Monster (2003), Theron portrayed serial killer Aileen Wuornos, a former prostitute who was executed in Florida in 2002 for killing six men (she was not tried for a seventh murder) in the late 1980s and early 1990s; film critic Roger Ebert felt that Theron gave "one of the greatest performances in the history of the cinema". For her portrayal, she was awarded the Academy Award for Best Actress at the 76th Academy Awards in February 2004, as well as the Screen Actors Guild Award and the Golden Globe Award. She is the first South African to win an Oscar for Best Actress. The Oscar win pushed her to The Hollywood Reporter's 2006 list of highest-paid actresses in Hollywood, earning up to US$10 million for a film; she ranked seventh. AskMen named her the number one most desirable woman of 2003.

For her role as Swedish actress and singer Britt Ekland in the 2004 HBO film The Life and Death of Peter Sellers, Theron garnered Golden Globe Award and Primetime Emmy Award nominations. In 2005, she portrayed Rita, the mentally challenged love interest of Michael Bluth (Jason Bateman), on the third season of Fox's television series Arrested Development, and starred in the financially unsuccessful science fiction thriller Æon Flux; for her voice-over work in the Aeon Flux video game, she received a Spike Video Game Award for Best Performance by a Human Female.

In the critically acclaimed drama North Country (2005), Theron played a single mother and an iron mine worker experiencing sexual harassment. David Rooney of Variety wrote: "The film represents a confident next step for lead Charlize Theron. Though the challenges of following a career-redefining Oscar role have stymied actresses, Theron segues from Monster to a performance in many ways more accomplished [...] The strength of both the performance and character anchor the film firmly in the tradition of other dramas about working-class women leading the fight over industrial workplace issues, such as Norma Rae or Silkwood." Roger Ebert echoed the same sentiment, calling her "an actress who has the beauty of a fashion model but has found resources within herself for these powerful roles about unglamorous women in the world of men." For her performance, she received Academy Award and Golden Globe Award nominations for Best Actress. Ms. magazine honoured her for this performance with a feature article in its Fall 2005 issue. On 30 September 2005, Theron received a star on the Hollywood Walk of Fame.

In 2007, Theron played a police detective in the critically acclaimed crime film In the Valley of Elah, and produced and starred as a reckless, slatternly mother in the drama film Sleepwalking, alongside Nick Stahl and AnnaSophia Robb. The Christian Science Monitor praised the latter film, commenting that "Despite its deficiencies, and the inadequate screen time allotted to Theron (who's quite good), Sleepwalking has a core of feeling". In 2008, Theron starred as a woman who faced a traumatic childhood in the drama The Burning Plain, directed by Guillermo Arriaga and opposite Jennifer Lawrence and Kim Basinger, and played the ex-wife of an alcoholic superhero alongside Will Smith in the superhero film Hancock. The Burning Plain found a limited release in US theatres, but grossed $5,267,917 outside the US. Hancock made US$624.3 million worldwide. Also in 2008, Theron was named the Hasty Pudding Theatricals Woman of the Year, and was asked to be a UN Messenger of Peace by the UN Secretary-General Ban Ki-moon.

Career hiatus and fluctuations (2009–2016)
Her film releases in 2009 were the post-apocalyptic drama The Road, in which she briefly appears in flashbacks, and the animated film Astro Boy, providing her voice for a character. On 4 December 2009, Theron co-presented the draw for the 2010 FIFA World Cup in Cape Town, South Africa, accompanied by several other celebrities of South African nationality or ancestry. During rehearsals she drew an Ireland ball instead of France as a joke at the expense of FIFA, referring to Thierry Henry's handball controversy in the play-off match between France and Ireland. The stunt alarmed FIFA enough for it to fear she might do it again in front of a live global audience.

Following a two-year hiatus from films, Theron returned to the spotlight in 2011 with the black comedy Young Adult. Directed by Jason Reitman, the film earned critical acclaim, particularly for her performance as a depressed divorced, alcoholic 37-year-old ghostwriter. Richard Roeper awarded the film an A grade, stating "Charlize Theron delivers one of the most impressive performances of the year". She was nominated for a Golden Globe Award and several other awards. Roger Ebert called her one of the best actors working today.

In 2012, Theron took on the role of villain in two big-budgeted films. She played Evil Queen Ravenna, Snow White's evil stepmother, in Snow White and the Huntsman, opposite Kristen Stewart and Chris Hemsworth, and appeared as a crew member with a hidden agenda in Ridley Scott's Prometheus. Mick LaSalle of the San Francisco Chronicle found Snow White and the Huntsman to be "[a] slow, boring film that has no charm and is highlighted only by a handful of special effects and Charlize Theron's truly evil queen", while The Hollywood Reporter writer Todd McCarthy, describing her role in Prometheus, asserted: "Theron is in ice goddess mode here, with the emphasis on ice [...] but perfect for the role all the same". Both films were major box office hits, grossing around US$400 million internationally each. The following year, Vulture/NYMag named her the 68th Most Valuable Star in Hollywood saying: "We're just happy that Theron can stay on the list in a year when she didn't come out with anything [...] any actress who's got that kind of skill, beauty, and ferocity ought to have a permanent place in Hollywood". On 10 May 2014, Theron hosted Saturday Night Live on NBC. In 2014, Theron took on the role of the wife of an infamous outlaw in the western comedy film A Million Ways to Die in the West, directed by Seth MacFarlane, which was met with mediocre reviews and moderate box office returns.

In 2015, Theron played the sole survivor of the massacre of her family in the film adaptation of the Gillian Flynn novel Dark Places, directed by Gilles Paquet-Brenner, in which she had a producer credit, and starred as Imperator Furiosa in Mad Max: Fury Road (2015), opposite Tom Hardy. Mad Max received widespread critical acclaim, with praise going towards Theron for the dominant nature taken by her character. The film made US$378.4 million worldwide. She next reprised her role as Queen Ravenna in the 2016 film The Huntsman: Winter's War, a sequel to Snow White and the Huntsman, which was a critical and commercial failure. In 2016, Theron starred as a physician and activist working in West Africa in the little-seen romantic drama The Last Face, with Sean Penn, provided her voice for the 3D stop-motion fantasy film Kubo and the Two Strings, and produced the independent drama Brain on Fire. That year, Time named her in the Time 100 list of the most influential people in the world.

Resurgence (2017–present)

In 2017, Theron starred in The Fate of the Furious as the cyberterrorist Cipher, the main antagonist of the entire franchise, and played a spy on the eve of the collapse of the Berlin Wall in 1989 in Atomic Blonde, an adaptation of the graphic novel The Coldest City, directed by David Leitch. The Fate of The Furious had a worldwide gross of US$1.2 billion. and Atomic Blonde was described by Richard Roeper of the Chicago Sun-Times as "a slick vehicle for the magnetic, badass charms of Charlize Theron, who is now officially an A-list action star on the strength of this film and Mad Max: Fury Road". In the black comedy Tully (2018), directed by Jason Reitman and written by Diablo Cody, Theron played an overwhelmed mother of three. The film was acclaimed by critics, who concluded it "delves into the modern parenthood experience with an admirably deft blend of humor and raw honesty, brought to life by an outstanding performance by Charlize Theron". She played the president of a pharmaceutical corporation in the crime film Gringo and produced the biographical war drama film A Private War, both released in 2018.

In 2019, Theron produced and starred in the romantic comedy film Long Shot, opposite Seth Rogen and directed by Jonathan Levine, portraying a U.S. Secretary of State who reconnects with a journalist she used to babysit. The film had its world premiere at South by Southwest in March 2019, and was released on 3 May 2019, to positive reviews from film critics. Theron next starred as Megyn Kelly in the drama Bombshell, which she co-produced. Directed by Jay Roach, the film revolves around the sexual harassment allegations made against Fox News CEO Roger Ailes by former female employees. For her work in the film, Theron was nominated for an Academy Award for Best Actress, Golden Globe Award for Best Actress in a Motion Picture – Drama, Critics' Choice Movie Award for Best Actress, Screen Actors Guild Award for Outstanding Performance by a Female Actor in a Leading Role, and BAFTA Award for Best Actress in a Leading Role. That same year, Forbes ranked her as the ninth highest-paid actress in the world, with an annual income of $23 million. In 2020, she produced and starred opposite KiKi Layne in The Old Guard, directed by Gina Prince-Bythewood. The following year, she reprised her role as Cipher in F9, originally set for release on 22 May 2020, before its delay to June 2021 due to the COVID-19 pandemic.

In 2022, Theron appeared in some projects. Upon the film's release in May, it was revealed that Theron would be portraying the character Clea in the Marvel Cinematic Universe (MCU), beginning with her debut in the mid-credits scene of the superhero film Doctor Strange in the Multiverse of Madness. She played Lady Lesso in the fantasy Netflix film The School for Good and Evil and filmed a cameo in Season 3 opener of The Boys as an actress playing Stormfront.

In 2023, Theron reprised her role as Cipher in Fast X and executive produced Last Call: When a Serial Killer Stalked Queer New York, a documentary series revolving around a serial killer preying upon gay men for HBO.

Activism

The Charlize Theron Africa Outreach Project (CTAOP) was created in 2007 by Theron, in an effort to support African youth in the fight against HIV/AIDS. By November 2017, CTAOP had raised more than $6.3 million to support African organizations working on the ground.

In 2008, Theron was named a United Nations Messenger of Peace. In his citation, Ban Ki-moon said of Theron: "You have consistently dedicated yourself to improving the lives of women and children in South Africa, and to preventing and stopping violence against women and girls." She recorded a public service announcement in 2014 as part of the UN's "Stop Rape Now" program.

In December 2009, CTAOP and Toms Shoes partnered to create a limited edition unisex shoe. The shoe was made from vegan materials and inspired by the African baobab tree, the silhouette of which was embroidered on blue and orange canvas. Ten thousand pairs were given to destitute children, and a portion of the proceeds went to CTAOP.

In 2020, CTAOP partnered with Parfums Christian Dior to create Dior Stands With Women, an initiative that includes Cara Delevingne, Yalitza Aparicio, Paloma Elsesser, and others, to encourage women to be assertive by documenting their journey, challenges and accomplishments.

Theron is involved in women's rights organizations and has marched in pro-choice rallies.

Theron is a supporter of same-sex marriage and attended a march and rally to support that in Fresno, California, on 30 May 2009. She publicly stated that she refused to get married until same sex marriage became legal in the United States, saying: "I don't want to get married because right now the institution of marriage feels very one-sided, and I want to live in a country where we all have equal rights. I think it would be exactly the same if we were married, but for me to go through that kind of ceremony, because I have so many friends who are gays and lesbians who would so badly want to get married, that I wouldn't be able to sleep with myself". Theron further elaborated on her stance in a June 2011 interview on Piers Morgan Tonight. She stated: "I do have a problem with the fact that our government hasn't stepped up enough to make this federal, to make [gay marriage] legal. I think everybody has that right".

In March 2014, CTAOP was among the charities that benefited from the annual Fame and Philanthropy fundraising event on the night of the 86th Academy Awards. Theron was an honoured guest along with Halle Berry and keynote speaker James Cameron.

In 2015, Theron signed an open letter which One Campaign had been collecting signatures for; the letter was addressed to Angela Merkel and Nkosazana Dlamini-Zuma, urging them to focus on women as they serve as the head of the G7 in Germany and the AU in South Africa respectively, which will start to set the priorities in development funding before a main UN summit in September 2015 that will establish new development goals for the generation. In August 2017, she visited South Africa with Trevor Noah and made a donation to the South African charity Life Choices. In 2018, she gave a speech about AIDS prevention at the 22nd International AIDS Conference in Amsterdam, organized by the International AIDS Society.

On 22 June 2022, it was announced that Theron and Sheryl Lee Ralph would receive the Elizabeth Taylor Commitment to End AIDS Award for their commitment to raising awareness of HIV at the Elizabeth Taylor Ball to End AIDS fundraising gala.

Endorsements
Having signed a deal with John Galliano in 2004, Theron replaced Estonian model Tiiu Kuik as the spokeswoman in the J'Adore advertisements by Christian Dior. In June 2024, Dior announced Theron will no longer be the face of J'Adore and be replaced by Barbadian singer and businesswoman Rihanna. In 2018, she appeared in a new advertisement for Dior J'adore. From October 2005 to December 2006, Theron earned US$3 million for the use of her image in a worldwide print media advertising campaign for Raymond Weil watches. In February 2006, she and her production company were sued by Weil for breach of contract. The lawsuit was settled on 4 November 2008. In 2018, Theron joined Brad Pitt, Daniel Wu and Adam Driver as brand ambassadors for Breitling, dubbed the Breitling Cinema Squad.

Personal life
In 2007, Theron became a naturalised citizen of the United States, while retaining her South African citizenship.

Theron has adopted two children: a daughter in March 2012 and another daughter in July 2015. She has been interested in adoption since childhood, when she became aware of orphanages and the overflowing numbers of children in them. In April 2019, Theron revealed that Jackson, then seven years old, is a transgender girl. She said of her daughters, "They were born who they are and exactly where in the world both of them get to find themselves as they grow up, and who they want to be, is not for me to decide".

She is inspired by actresses Susan Sarandon and Sigourney Weaver. She has described her admiration for Tom Hanks as a "love affair" and watched many of his films throughout her youth. Hollywood actors were not featured in magazines in South Africa so she did not know how famous he was until she moved to the United States, which has been inferred as a factor of her "down-to-earth" attitude to fame. After filming for That Thing You Do! finished, Theron got Hanks' autograph on her script. She later presented him his Cecil B. DeMille Award in 2020, in which Hanks revealed that he had a mutual admiration for Theron's career since the day he met her.

Theron said in 2018 that she went to therapy in her thirties because of anger, discovering that it was due to her frustration growing up during South Africa's apartheid, which ended when she was 15.

In 2019, Theron spoke about her method of working on roles. Creating a physical identity together with the emotional part of the character, she said, is "a great tool set that adds on to everything else you were already doing as an actor. It's a case-by-case thing, but there is, to me, this beautiful thing that happens when you can get both sides: the exterior and interior. It's a really powerful dynamic". When preparing for a role, "I almost treat it like studying. I will find space where I am alone, where I can be focused, where there's nobody in my house, and I can really just sit down and study and play and look at my face and hear my voice and walk around and be a fucking idiot and my dogs are the only ones who are seeing that".

Theron is a longtime fan of the English band Depeche Mode, and was the presenter for their Rock and Roll Hall of Fame induction in 2020.

Relationships
Theron's first public relationship was with actor Craig Bierko, whom she dated from 1995 to 1997.

Theron was in a three-year relationship with singer Stephan Jenkins until October 2001. Some of Third Eye Blind's third album, Out of the Vein, explores the emotions Jenkins experienced as a result of their breakup.

Theron began a relationship with Irish actor Stuart Townsend in 2001 after meeting him on the set of Trapped. The couple lived together in Los Angeles and Ireland. The couple split up in late 2009.

In December 2013, Theron began dating American actor Sean Penn. The relationship ended in June 2015. There were reports they were engaged, and Theron said that they were never engaged. Theron starred in Penn's film The Last Face (2016), which they filmed while still a couple.

Health concerns
Theron often quips that she has more injuries on sets that are not action films; however, while filming Æon Flux in Berlin, Theron suffered a herniated disc in her neck, caused by a fall while filming a series of back handsprings. It required her to wear a neck brace for a month. Her thumb ligament tore during filming of The Old Guard when her thumb caught in another actor's jacket during a fight scene, which required three operations and six months in a thumb brace. During the filming of Atomic Blonde she broke teeth from clenching her jaw, and had dental surgery to remove them: "I had the removal and I had to put a donor bone in there to heal until I came back, and then I had another surgery to put a metal screw in there."

Outside of action films, she had a herniated disk in her lower back as she filmed Tully and suffered from a depression-like state, which she theorized was the result of the processed food she had to eat for her character's post-natal body. In July 2009, she was diagnosed with a serious stomach virus, thought to be contracted while overseas. While filming The Road, Theron injured her vocal cords while screaming during the labour scenes. When promoting Long Shot, she revealed that she laughed so hard at Borat that her neck locked for five days. She added that on the set of Long Shot she "ended up in the ER" after knocking her head against a bench behind her when she was putting on knee pads.

Filmography and accolades
Theron has appeared in over 50 films. She has received an Academy Award, a Golden Globe Award, and nominations for three British Academy Film Awards and an Emmy Award.

license: Creative Commons - Attribution Share-Alike - https://creativecommons.org/licenses/by-sa/4.0/

title: Charlize Theron