While their population centers were

While their population centers were along the coast, the settlements were growing into the interior.

Resultaten (Nederlands) 1:

Terwijl hun bevolking centra langs de kust waren, groeiden de nederzettingen in het interieur.

Resultaten (Nederlands) 2:

Terwijl hun bevolkingscentra waren langs de kust werden de nederzettingen groeien in het interieur.

Resultaten (Nederlands) 3:

Hoewel ze in de centra van de bevolking langs de kust van nederzettingen is toegenomen.