I would never dream of sharpening an image until it is at the final resolution. I need different levels for a 300x300, 500x500, 1000x1000, 1500x1500, and larger images. The smaller the image the more sharpening is needed and what would be appropriate for a 2400 x 3000 image is not going to be sufficient for a smaller image.
I reduce the images to the size needed for the layouts, sharpen, and save as JPEGs and then bring them into the page layouts. I know in advance the sizes I want to use for each image based on its value in the layout. The higher the value the larger the image and the less the sharpening that is applied.
I work the layouts around the subsets of images. One set may have 3 small vertical images and a large horizontal and the next spread may have 2 large verticals, and so forth. I create the layouts based on these sets of images. I realize it is a different approach that is not common for nature photography but it is very effective nonetheless.