Summary: The Lowlands have lived in peace ever since the Revolution. But now a new threat emerges. They outnumber any army and they have a deadly art. Sucked into all of this is Che Maker, Totho, Tynisa, and Salma. They all want to help Che's uncle and save the Lowlands. Do they have what it takes?
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The novel opens with a prologue set in the city of Myna. The city is under siege by the aggressive and expansionistic Wasp Empire and a group of students and activists from the Great College of Collegium have come to witness the Wasp threat for themselves. Stenwold Maker, a Beetle-kinden man, observes through a telescope as Wasp automotives ram the city's gates and is dismayed when an explosive trap he had helped set there fails to detonate. One of his group, a Spider-kinden woman Atryssa, has failed to turn up and the trap's failure implies she has betrayed them. With the city quickly falling Stenwold and his Ant-kinden friend Marius are joined by Tisamon, a Mantis-kinden and the last member of their group, and they fall back to Myna's highest airfield, their retreat shielded by a squad of Ant-kinden soldiers led by Marius. Tisamon had overcome his race's deep-seated hatred of Spider-kinden to have a love affair with Atryssa and her apparent betrayal sends him into a suicidal killing frenzy fuelled by grief and anger, and Stenwold barely manages to talk him into boarding the aircraft he has secured. Stenwold kills his first man, a Wasp soldier who attacked him, while most of Marius's squad die defending the airfield. Only Marius and a single female Ant make it to the aircraft, and Marius mortally wounded by a crossbow bolt. Stenwold pilots the aircraft away while Myna falls to the Wasps, Marius dies and Tisamon seethes.
The story picks up twelve years later in Collegium. Stenwold is now a Master of the Great College teaching contemporary history and considered a radical for his actions over the last twelve years trying to open everyone's eyes to the Wasp threat. He has become a spy master, cultivating agents from among the College's students and sending them out into the Lowlands to discover Wasp activities. The Wasp Empire itself has been occupied with a Twelve-Year War against the Dragonfly Commonweal to the north of the Lowlands but is now turning its eyes back west and Stenwold is the only one who acknowledges the threat. He has cultivated two new agents, his foster-daughter Tynisa , a Spider-kinden, and Salme Dien , a Dragonfly-kinden nobelman's son sent to the Lowands to evaluate the Wasp threat. Together with Stenwold's niece Cheerwell Maker and an Ant-Beetle halfbreed artificer student named Totho they make up a duelling team dubbed 'the Magnificient Felbling.' Tynisa and Salma use the duels as practice for their lives as Stenwold's agents.
After an attempt on Stenwold's life by thugs hired by the Wasps he decides to send Tynisa and Salma on the airship Sky Without to the industrial city of Helleron to meet with his associates there. Totho also makes an impression on Stenwold and he includes him in the group, but it is only after the attempted assassination that Stenwold agrees to let Cheerwell go as well, taking his place on the airship so his presence will not endanger the young agents. Once aboard the airship and bound for Helleron Tynisa seduces a Wasp captain on the ship, Halrad, to keep him from investigating them too hard. However, the tables turn when Thalric, a Rekef Outlander agent for the Wasp Empire tasked with ending Stenwold's activities, arrives on board looking for him. Once he discovers them the agents are forced to flee the ship, stealing the flier he arrived in and killing some of their pursuers. They crash-land a few miles from Helleron and are forced to complete their journey on foot, arriving later than planned. By this time the man they were meant to meet with, Bolwyn, has been killed by Scyla, a shape-shifting Spider-kinden working with Thalric, and their rendezvous is a trap. Though all four agents escape they become seperated. Totho is rescued by Scuto, a Thorn-bug kinden and Stenwold's head agent in Helleron who controls his own espionage operation within the city. Tynisa flees through the streets, increasingly taken over by her killing instincts, and ends up among a street gang who her take her in as an enforcer. Cheerwell and Salma make their way to one of Cheerwell's relatives within the city, Elias Monger. While staying at his house outside the city Cheerwell encounters a young Moth-kinden man, Achaeos, who was injured in a raid on Monger's mine workings. Rather than turn him in she overcomes his mistrust and tends to his wound.
Tynisa becomes involved in her adopted gang's territorial feuds and witnesses some of her boss's men wiped out by a Mantis-kinden mercenary, Tisamon. Later she manages the leave the gang after doing good service for them and makes her way to Scuto's hideout, where she is reunited with Totho. Stenwold, having heard what befell the group, travels to Helleron himself and arrives at Scuto's, relived to find Tynisa and Totho safe. However by this point Elias Monger has turned Cheerwell and Salma over to Thalric, who has taken them prisoner, unknowingly witnesses by Achaeos. Stenwold meets with Tisamon for the first time in nearly a decade a recruits him once more. Together they go to Monger's house, where Wasp soldiers lie in wait. However, Achaeos warns Tisamon who is able to save Stenwold from capture and kill the Wasps. Stenwold confronts Monger for selling out Salma and his niece but holds back from killing him because he does not wish to kill family. Tisamon, however, is not related to Monger and Stenwold allows him to kill him.
Thalric takes Cheerwell and Salma as prisoners to Asta, formerly a small desert trading town turned staging camp for the Wasp Empire's armies. Cheerwell is almost raped by a Wasp slaver named Brutan but Brutan holds back at the last second out of fear of Thalric. Unconcerned, Thalric questions Cheerwell about her uncle before recieving summons to Myna to investigate his old mentor Ulther for corruption. He travels on, taking Cheerwell and Salma with him.
Stenwold, Tynisa, Totho, Tisamon and Achaeos travel towards Asta to try and rescue Cheerwell and Salma while Scuto remains in Helleron. Stenwold does something he has not had the oportunity or courage to do before now, introducing Tynisa to Tisamon. Tisamon, recognizing Tynisa as almost identical to Atryssa, threatens to kill Stenwold unless he explains. Stenwold tells him that Atryssa never betrayed them at Myna, but was discovered by Wasp agents and forced to flee. She gave birth to Tynisa, her and Tisamon's daughter, before dying, entrusting Tynisa to Stenwold. Though she appears full-blood Spider, Tynisa is in fact an unheard-of Spider-Mantis halfbreed.
The group arrive an the outskirts of Asta, near the haunted Darakyon forest. While wary of one another, Tynisa and Tisamon are the best scouts and they infiltrate Asta to search for Cheerwell and Salma. While they are gone Stenwold, Totho and Achaeos are chanced upon by a Wasp patrol and forced to flee into the Darakyon. Pursued by nameless terrors that kill the Wasps chasing them Stenwold and Totho manage to escape the forest and return to their camp, but Achaeos gets seperated and confronted by the evil, twisted spirits that dwell in the woods. He flees, scarred by the experience, and finds the other two some time after their own escape, arousing Totho's suspicions. Totho has had feelings for Cheerwell since their first meeting and mislikes and distrusts Achaeos, seeing him as a rival for her. Tisamon and Tynisa return having acted as one during the search but found nothing, and Tisamon reverts to ignoring Tynisa which frustrates her.
Cheerwell and Salma are impriosned in Myna while Thalric begins his mission for the Rekef to investigate Ulther, the city's Wasp governor. Thalric straps Cheerwell to a torture table, apparently about to forcibly draw information on Stenwold from her, but only uses the noise made by the torture machine to conduct a clandestine conversation with his ally Aagen about exposing Ulther. In the end he forces matters with his former mentor quite simply, confiscating a rare and beautiful Butterfly-kinden woman, Grief in Chains, from Ulther's harem. Ulther sends men to kill Thalric but Thalric survives the fight and goes after Ulther.
Meanwhile Stenwold, Tisamon, Totho, Tynisa and Achaeos have arrived in Myna and made contact with the anti-Wasp resistance there. The resistance allies with them for a raid on the Governor's Palace, with the twin goals of freeing Cheerwell and Salma but also Kymene, the resistance leader Ulther has captured. Their raid coincides with Thalric's move on Ulther and once Kymene is freed she and her resistance followers allow Thalric to carry out his mission as it will destabilize the Wasp Empire's control of Myna. Thalric kills Ulther while the resistance and Stenwold's group escape, having freed Cheerwell and Salma. In the tunnels beneath the city Tynisa hangs back and confronts Tisamon, engaging him in a deadly duel to force him to acknowledge her as his daughter. Tisamon eventually beats her but she gave him a challenge no-one had for years and he acknowledges her.
Stenwold's group pull out of Myna. By this point Cheerwell and Achaeos have fallen in love and he takes her to his mountaintop city of Tharn where he has become an outsider among his people. They sleep together, which unlocks Cheerwell's Art, enabling her to fly. Salma, who has become infatuated with Grief in Chains since encountering her during his capture, learns that she is among the healers following the Wasp army that is descending on the Ant-kinden city of Tark in the Lowlands. He leaves the group, following a resistance guide to Tark to find his love.
Having seen Cheerwell and Achaeos in love and unable to bear it Totho departs the group in the night, leaving to follow Salma to Tark and unknowingly allowing the shape-shifter Scyla, who has been tailing them since Helleron, to assume his identity. After Stenwold's group arrive at Helleron Scyla, as Totho, betrays the location of Scuto's hideout to the Wasps before following Cheerwell and Achaeos. Achaeos sees through her disguise and badly wounds her with an arrow, making Cheerwell turn on him in rage until she realizes what happened. They return to Helleron, ousted by the Moths of Tharn, and re-join Stenwold and the others. In the city Stenwold and Thalric encounter one another and explore their mutual grudging respect over a drink. From their conversation Stenwold pieces together Thalric's next move and rushes back to Scuto's hideout which has come under attack. Warned just in time by a member of Tynisa's former gang Stenwold and Scuto's groups defend the hideout from the surprise assault but several of Scuto's people are killed and they are forced to flee. Enough of them remain however to take action when Stenwold works out that Thalric and his people are going to seize the steam-train Pride and use it to spearhead a surprise attack on Collegium. Joined by some Moths from Tharn who were swayed by Achaeos the rag-tag group defends the Pride in a fierce skirmish. Thalric badly wounds Tisamon with his sting but is badly wounded in turn by Tynisa and Scuto succeeds in blowing up the train before the Wasps can use it. The Moths return to Tharn and Stenwold, Cheerwell, Tisamon, Tynisa, Achaeos, Scuto and his few surviving agents return to Collegium.