middle Motagua River drainage, due to the arid climate. It covered a broad area that included Cubulco, Rabinal, and Salamá (all in Baja Verapaz), San Agustín de la Real Corona (modern San Agustín Acasaguastlán) and La Magdalena in El Progreso, and Chimalapa, Gualán, Usumatlán and Zacapa, all in the department of Zacapa. Chimalapa, Gualán and Usumatlán were all satellite settlements of Acasaguastlán. San Cristóbal Acasaguastlán and the surrounding area were reduced into colonial settlements by friars of the Dominican Order; at the time of the conquest the area was inhabited by Poqomchiʼ Maya and by the Nahuatl-speaking Pipil. In the 1520s, immediately after conquest, the inhabitants paid taxes to the Spanish Crown in the form of cacao, textiles, gold, silver and slaves. Within a few decades taxes were instead paid in beans, cotton and maize. Acasaguastlán was first given in encomienda to conquistador Diego Salvatierra in 1526. Chiquimula Chiquimula de la Sierra ("Chiquimula in the Highlands"), occupying the area of the modern department of Chiquimula to the east of the Poqomam and Chajoma, was inhabited by Chʼortiʼ Maya at the time of the conquest. The first Spanish reconnaissance of this region took place in 1524 by an expedition that included Hernando de Chávez, Juan Durán, Bartolomé Becerra and Cristóbal Salvatierra, amongst others. In 1526 three Spanish captains, Juan Pérez Dardón, Sancho de Barahona and Bartolomé Becerra, invaded Chiquimula on the orders of Pedro de Alvarado. The indigenous population soon rebelled against excessive Spanish demands, but the rebellion was quickly put down in April 1530. However, the region was not considered fully conquered until a campaign by Jorge de Bocanegra in 1531–1532 that also took in parts of Jalapa. The afflictions of Old World diseases, war and overwork in the mines and encomiendas took a heavy toll on the inhabitants of eastern Guatemala, to the extent that indigenous population levels never recovered to their pre-conquest levels. Campaigns in the Cuchumatanes In the ten years after the fall of Zaculeu various Spanish expeditions crossed into the Sierra de los Cuchumatanes and engaged in the gradual and complex conquest of the Chuj and Qʼanjobʼal. The Spanish were attracted to the region in the hope of extracting gold, silver and other riches from the mountains but their remoteness, the difficult terrain and relatively low population made their conquest and exploitation extremely difficult. The population of the Cuchumatanes is estimated to have been 260,000 before