Full context:   The proportions of the human hand are plesiomorphic (shared by both ancestors and extant primate species); the elongated thumbs and short hands more closely resemble the hand proportions of Miocene apes than those of extant primates. Humans did not evolve from knuckle-walking apes, and chimpanzees and gorillas independently acquired elongated metacarpals as part of their adaptation to their modes of locomotion. Several primitive hand features most likely present in the chimpanzee–human last common ancestor (CHLCA) and absent in modern humans are still present in the hands of Australopithecus, Paranthropus, and Homo floresiensis. This suggests that the derived changes in modern humans and Neanderthals did not evolve until 2.5 to 1.5 million years ago or after the appearance of the earliest Acheulian stone tools, and that these changes are associated with tool-related tasks beyond those observed in other hominins. The thumbs of Ardipithecus ramidus, an early hominin, are almost as robust as in humans, so this may be a primitive trait, while the palms of other extant higher primates are elongated to the extent that some of the thumb's original function has been lost (most notably in highly arboreal primates such as the spider monkey). In humans, the big toe is thus more derived than the thumb.