has also driven vicariant speciation in Macowania daisies in South Africa's Drakensberg mountains, along with Dendrocincla woodcreepers in the South American Andes. The Laramide orogeny during the Late Cretaceous even caused vicariant speciation and radiations of dinosaurs in North America. Adaptive radiation, like the Galapagos finches observed by Charles Darwin, is often a consequence of rapid allopatric speciation among populations. However, in the case of the finches of the Galapagos, among other island radiations such as the honeycreepers of Hawaii represent cases of limited geographic separation and were likely driven by ecological speciation. Isthmus of Panama Geological evidence supports the final closure of the isthmus of Panama approximately 2.7 to 3.5 mya, with some evidence suggesting an earlier transient bridge existing between 13 and 15 mya. Recent evidence increasingly points towards an older and more complex emergence of the Isthmus, with fossil and extant species dispersal (part of the American biotic interchange) occurring in three major pulses, to and from North and South America. Further, the changes in terrestrial biotic distributions of both continents such as with Eciton army ants supports an earlier bridge or a series of bridges. Regardless of the exact timing of the isthmus closer, biologists can study the species on the Pacific and Caribbean sides in what has been called, "one of the greatest natural experiments in evolution". Additionally, as with most geologic events, the closure was unlikely to have occurred rapidly, but instead dynamically—a gradual shallowing of sea water over millions of years. Studies of snapping shrimp in the genus Alpheus have provided direct evidence of an allopatric speciation event, as phylogenetic reconstructions support the relationships of 15 pairs of sister species of Alpheus, each pair divided across the isthmus and molecular clock dating supports their separation between 3 and 15 million years ago. Recently diverged species live in shallow mangrove waters while older diverged species live in deeper water, correlating with a gradual closure of the isthmus. Support for an allopatric divergence also comes from laboratory experiments on the species pairs showing nearly complete reproductive isolation. Similar patterns of relatedness and distribution across the Pacific and Atlantic sides have been found in other species pairs such as: Diadema antillarum and Diadema mexicanum Echinometra lucunter and Echinometra vanbrunti Echinometra viridis and E. vanbrunti Bathygobius soporator and Bathygobius ramosus B. soporator and Bathygobius andrei Excirolana braziliensis and variant morphs Refugia Ice ages have played important roles