The Nicoya Peninsula and central Pacific coastline were settled by the indigenous Matambú people around 500 AD. Escaping slavery from southern Mexico, they were drawn to the area by the wealth of the ocean and fertile hills. Spanish conquistadors explored the region in the early 16th century and established short-lived settlements. These fell victim to tropical disease and the ferocious resistance of the Matambú. However, the indigenous population was ultimately overwhelmed by Spanish forces. Founded in 1522, Puntarenas was the principal city of the region by the early 1800s. It flourished due to the 19th century coffee trade, and developed into the nation’s main port for coffee exports to Europe. In the early decades of the 20th century, bananas were planted along the narrow coastal plain farther south. They were replaced in the 1970s by African oil palms, which extend for miles between the shore and forested mountains. Much of the region, notably Southern Nicoya, was heavily denuded during the last century, but major conservation and reforestation efforts are now extending the protected areas.