About 3 million years ago, the Isthmus of Panama (a narrow strip of land connecting North and South America) formed, dividing marine organisms into Pacific and Caribbean populations. Researchers have examined species of snapping shrimp on both sides of the isthmus. Based on morphology, there appear to be seven pairs of species, with one species of each pair in the Pacific and the other in the Caribbean. Researchers reconstructed a phylogeny using mitochondrial DNA sequences and found that each of the species-pairs, separated by the isthmus, were indeed each other's closest relatives. The researchers also found that species from many pairs were not very interested in courting with each other under lab conditions, and any that did mate almost never produced fertile offspring. The species pairs on opposite sides of the isthmus are true species under which species concept(s)?