Darla was born under a different name in the late 16th century in the British Isles. As a young prostitute, she immigrated to the Virginia Colony in North America and became independently wealthy, but also contracted a fatal case of syphilis. By 1609, the prostitute was dying in her luxurious house. Because of her dislike of religion, she scoffed at a priest who came to her deathbed, despite having called for one the previous night in her delirium. When asked to beg for God's forgiveness, she claimed her soul was well past saving and that the Devil may take it. The priest then revealed his true identity: The Master, leader of the elite cult of vampires known as the Order of Aurelius. Believing he was death, she was corrected by the Master who claimed he was actually her savior. He sired her and at some point gave her the name "Darla" (Gaelic for "dear one") and she eventually forgot her original name. As a vampire, Darla developed a taste for luring male victims by appearing as a seductive and mysterious yet harmless woman.

By 1753, Darla was out exploring the world and wreaking havoc, during which time she entered a village in Galway, Ireland. There she found a handsome young man named Liam, who was drinking and whoring his way through a rather uneventful life. Admiring him from afar, Darla lured Liam into an alley where she seduced him by offering to show him things he had never imagined. She sired Liam and chose to make him her mate, naming him Angelus, inspired by his sister's belief that he was an angel come back to her.

In 1760, after cutting a bloody swath across Wales and England, Darla brought her new consort back to The Master in London. However, Angelus had no interest in the Order of Aurelius and its tenets of underground dwelling and tribute to the Old Ones. Being told to choose between her sire and the pleasures Angelus offered her, Darla chose her lover and left with him.

In 1764, Angelus and Darla were back in York, where they were hunted down by Daniel Holtz, a vampire hunter. The lovers decided to punish Holtz by luring him to an empty cottage, leaving behind the message "How do you hope to save others, when you can't save your own?" While Holtz was on this chase, Angelus and Darla raped and killed Holtz's wife, Darla sired his daughter, and they killed his infant son as well. After being forced to slay his own daughter, Holtz swore revenge, and began a crusade to eliminate the two vampires that would last nearly an entire decade.

By 1765, Darla and Angelus fled from Italy to France with Holtz hot on their heels. Holtz managed to trap the pair within a burning barn, at which point Darla hit Angelus with a shovel and escaped on the only horse left. Angelus followed Darla all the way to Vienna, where he made her "pay for [Darla's] sins. Again and again".

In 1767, Darla and Angelus joined a couple of young vampire lovers, James and Elisabeth, in Marseilles. The couples burned the village of Comte de Leon to the ground, which attracted the attention of Holtz. While Darla and Elisabeth escaped to the ship that would take the four vampires to Morocco, James and Angelus stayed behind to face the vampire hunter. But resentful of Darla's betrayal in the barn, Angelus revealed his lover's whereabouts to save his own life.

Despite Angelus' betrayal, Darla remained with Angelus as the two traveled through North Africa trying to escape Holtz. The pair were in Rome when the relentless Holtz found them in 1771. Holtz managed to capture Angelus with the aid of a traditionalist Catholic sect known as Inquisitore, although with the aid of a group of local vampires, Darla saved her lover from Holtz's torture. Angelus wondered if they should kill Holtz, but Darla answered that it was much more fun to further ruin his life.

Back in London, the "family" of Darla and Angelus grew in 1860, when Darla showed him Drusilla, a young, pious girl blessed with visions, whom Angelus stalked and tortured into insanity before siring her. The trio remained unchanged until 1880, the year in which Darla told Drusilla to turn the "first drooling idiot that comes along." Drusilla then turned William, a romantic, heart-broken poet. Around the same time William was turned, Darla had a fight with Angelus and returned to the side of the Master. Nonetheless, the couple reconciled shortly thereafter, only to be forced to leave London because William's attacks were drawing unnecessary attention. The foursome fled all the way to Yorkshire where they were forced to hide in abandoned mines, much to Darla's dismay. Around this time, Darla referred to herself, Angelus, Drusilla, and Spike as The Whirlwind.

By 1894, the foursome had traveled to Rome where they crossed paths with The Immortal, a mysterious being who had already met the vampires in Frankfurt and saved a group of nuns from Angelus. While Angelus and Spike were kidnapped on the Immortal's orders, he seduced Darla and Drusilla and "violated" the two willing vampires in a m�nage � trois, a privilege they had never allowed Spike or Angel. A few years later, the four vampires traveled to Budapest to have some fun during the chaos of an earthquake.

In 1898, the Whirlwind traveled to Borsa, Romania. There, Darla gave Angelus a gift that would alter their lives forever--the favorite daughter of Clan Kalderash, a tribe of Roma. Angelus had his way with the girl for a night while Darla watched. His actions incurred the wrath of the Kalderash Elders, however, and one of their women performed a curse to restore Angelus' human soul. Angelus sought Darla's help, only to be rejected for having a "filthy soul."

Nevertheless, Darla took Spike and Drusilla to massacre the Kalderash. As the two lovers slaughtered everyone they found, Darla tried to reason with the father of the Kalderash daughter and convince him to have Angelus' soul removed in exchange for sparing the lives of his wife and daughters. Darla's leverage was lost when they saw that Spike had already killed the Elder's family. Without further hope, Darla snapped the Elder's neck and ordered the two to massacre the entire tribe.

In November 1898, Darla and Drusilla almost ended up being Brides of Dracula, although the famous vampire escorted them to Whitby, Yorkshire with the intention of having them stake each other. Spike then led a mob to Dracula's castle, and during the assault, fled with Darla and Drusilla.

Darla, who had developed a taste for religious wars, took Spike and Drusilla to China in 1900 where the trio found themselves in the midst of the Boxer Rebellion. Angelus, who had followed them, begged Darla to accept him again, hoping to prove to his former lover and himself that he could still be part of The Whirlwind. However, Angelus limited himself to killing thieves, murderers, and rapists. He even tried to save a missionary family. Darla discovered the family�a father, mother, daughter, and infant child�killing all but the baby, whom she kept as a test for Angelus. Incapable of killing an innocent, Angelus fled with the baby, forever renouncing Darla.

In 1901, Darla and "Angelus" cross paths for the last time in Budapest. By October 1934, Darla was living in Berlin, Germany. In September 1943, Darla had moved to England. By 1959, Darla had moved to the Hellmouth, Sunnydale, California with The Master.

After so many centuries, Darla still lured her victims by pretending to be a timid, innocent young woman, even adding a schoolgirl uniform to her act. One of those she killed was an ex-student that she drained on the grounds of Sunnydale High. It was the inadvertent warning sign for Buffy Summers (the Slayer) that vampires were active in Sunnydale.

Darla came face to face with Buffy the night prior to the Harvest, when she seduced Sunnydale High student Jesse McNally. Even though Jesse was intended to be an offering to the Master, Darla bit Jesse, for which she would be reprimanded. Despite facing Buffy, however, Darla managed to take Jesse underground thanks to the intervention of Luke, the Master's leading acolyte. The following night, Darla joined Luke, the newly sired Jesse, and other vampires in their attack against The Bronze, where Luke intended to perform the Harvest in order to release the Master from his mystical prison. While Luke, Jesse, and other vampires were dusted by Buffy and the Scooby Gang, Darla was able to escape after Willow Rosenberg threw holy water at her.

For weeks, Darla laid low while several members of the Order fell to Buffy's stake. She offered to kill the Slayer as an offering for her sire, but he sent The Three, a trio of ancient vampire warriors, instead. However, the Three failed due to the intervention of Darla's former lover Angelus, who had renamed himself "Angel" and allied himself with the Slayer. As penance, the Three offered their lives to the Master, and he allowed Darla to dust them, a task she gladly completed.

Understanding that brute force wouldn't be enough to kill Buffy, Darla devised a plan to turn her against Angel, and force one to kill the other, though she hoped Angel would kill Buffy in self-defense and, having experienced killing once again, return to her side. Darla pretended to be a classmate of Buffy's in order to be invited to the Summers residence. Once inside, she attacked Buffy's mother, biting her and leaving her in Angel's arms. As Darla had planned, Buffy arrived to see her injured mother held by Angel in vampire face, and concluded that Angel was drinking from her mother. The vampire and the Slayer faced each other in the Bronze, but Buffy eventually realized that Angel was not responsible. Darla, carrying a pair of pistols with her, arrived to ensure Buffy would die. However, Darla's plan turned against her, as Angel staked her in order to protect the woman he loved.

Nearly three years later, the evil law firm Wolfram & Hart performed the Raising ritual from the Scrolls of Aberjian to resurrect Darla as a human to torment Angel and/or to turn him evil. During her short time as a human, Darla found it increasingly difficult to live with a soul and desperately wanted to be a vampire again. She also seemed to become close to Wolfram & Hart lawyer Lindsey McDonald, though whether this was a ploy or not is uncertain. Soon she learned that she was dying of the disease that had almost killed her before she was sired; syphilis. Though it was now easily cured when diagnosed early enough, Wolfram & Hart was, as Lindsey said, about a month and 400 years too late. It made her even more desperate to be turned into a vampire, but instead, Angel, through acts of bravery and nobility, finally convinced her to live out her remaining days as human. Angel then went through the trials to win Darla a new chance at life. Though he endured, it was for naught as Darla had already gotten her second chance at life, and Angel reassured her that he would be there for her. Once she had fully accepted and embraced her humanity and soul, Wolfram & Hart, angry that their plans were going awry, brought in Drusilla to sire Darla.

After Darla rose again, she was initially upset over becoming a vampire again and attacked Drusilla. However, she quickly embraced it once more. She and Dru then celebrate Darla's raising by attending a Wolfram & Hart function at the home of Holland Manners. Darla and Drusilla massacred most of the staff attending the party out of revenge for being used as pawns. The only two survivors were Lindsey McDonald and Lilah Morgan, who the two vampires wanted to use as means to get what they want. Afterward, Darla and Drusilla attempted to recruit a gang of demons in order to take over Los Angeles, though they were all slaughtered by Angel. Shortly afterwards, Angel, who had descended into darkness, set both women on fire, though they did not die but were severely injured. As Drusilla went to Sunnydale to unsuccessfully try to bring Spike back into the fold, Darla hid in Lindsey's apartment to recover and hide from Wolfram & Hart. It's possible she also exploited Lindsey's romantic interest in her.

Angel's life perspective grew even darker when Holland Manners informed him that Wolfram & Hart's power came from the evil within humanity itself and that none of his good deeds would ever make a difference. Returning home, he realized Darla was laying in wait for him. He roughed her up and they had sex, with Darla gleefully expecting him to lose his soul, as the curse breaks when Angel experiences a moment of perfect happiness such as when he made love to Buffy Summers. The next morning, Angel still had his soul; sex with Darla made him experience perfect despair, rather than perfect happiness. Apologizing to Darla for his 'failure' to save her, Angel admitted that a part of him had seen it as part of his efforts to save himself�since she had created him, he had hoped that redeeming her would prove him worthy of redemption�but he now recognized that she was nothing to him, unable to even hate her. Surprised, angry, and upset, Darla attempted to stake Angel after he rejected her offer of a "repeat performance" to "get it right", only for Angel to grab the stake, informing Darla that he would allow her to live for now in gratitude for what she had done for him, but vowing that he would kill her next time he met her. As a result, Darla left Los Angeles.

However, the unthinkable had happened: Darla had become pregnant as a result of Angel winning a life by enduring the trials in an attempt to save her when she was dying. Darla visited every shaman in the Western Hemisphere, all of whom told her that her pregnancy was inherently impossible, but also impossible to abort. With nowhere else to turn, Darla went to Angel. Various theories emerged about the child; from a cult of vampires who worshiped it as a miracle, to the fear that it was the evil spoken of in the Tro-Clon prophesies. It turned out the child was simply a healthy human baby with whom the pregnant Darla was sharing a soul, allowing her to feel genuine love for the child and prompting her to seek innocent blood to feed it. This did not matter to Daniel Holtz, who had been brought through time by the demon Sahjhan to kill Angel and Darla. Darla realized that she would lose her soul after giving birth and would be unable to love her child. She feared that she would kill the child eventually as her vampiric nature re-asserted itself once the child's soul was gone from her body. Additionally, her undead body was becoming increasingly unable to sustain the life of her unborn child in time for it to be born.

When Darla went into labor, things did not look well, and it appeared that she would lose the baby. Adding to the difficulties of birth, Darla and Angel were being pursued by the murderous Holtz, who chased them into the alley behind Caritas during a violent rainstorm. Darla realized that the baby would die in the alley, just as the human Liam had died in an alley in order to birth the vampire Angelus. After telling Angel to make sure that their son would know that he was the one good thing the two vampires had ever done together, Darla staked herself, leaving behind a pile of dust and a crying and healthy infant�a final act of redemption to ensure the life of their son, Connor.

Sometime later, Darla's spirit appeared to a teenaged Connor when he was being tempted by Cordelia Chase (possessed by the higher power later known as Jasmine) to murder an innocent teenage girl so that her blood could anoint a pregnant Cordelia so she could give birth to Jasmine. Sent by The Powers That Be, Darla pleaded with her son to not commit murder. Seemingly moved by his mother's words, Connor began to untie the girl, but "Cordelia" chastised him for allowing himself to be swayed by Darla's spirit, whose presence she could sense. "Cordelia" and Darla (the former clad completely in black and the latter in white) competed for Connor's soul, with Darla telling Connor to listen to his heart and "Cordelia" telling him that Darla was just an illusion created by Angel's team using magic in order to trick him. Cordelia's words combined with Connor's anxiety for the life of the "baby," ultimately won out and Connor exclaimed to Darla, "You are not my mother!" and dragged the terrified innocent girl to "Cordelia." Connor saw the girl take on Darla's form for a moment though "Cordelia" promptly killed her. The girl's corpse remained as Darla for a brief moment. Connor never mentioned what he saw to anybody.

When Spike went up against Wolfram & Hart in Las Vegas, the liaison to the Senior Partners, possessing human Jeremy Johns, told him that they were planning on creating a fake Darla in order to get to Connor.