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"His topknot is like a crown." (Secondary characteristics, No 53)
"He has a topknot as if crowned with a flower garland." (Secondary characteristics, No 80)
The first representations of the Buddha in the 1st century CE in the Greco-Buddhist art of Gandhara also represent him with a topknot, rather than just a cranial knob. It is thought that the interpretation of the ushnisha as a supernatural cranial protuberance happened at a later date, as the representation of the topknot became more symbolic and its original meaning was lost.
The Boddhisattva-Cakravartin in Early Buddhism
In Early Buddhism, the uṣṇīṣa was represented differently. The Mahāvastu (1.259f) and the Divyāvadāna, as well as the Theravadin Milindapañha, describe the marks of the cakravartin, an idealised world-ruler: uṣṇīṣa or patka turban, chhatra parasol, "horn jewel" or vajra, whisk and sandals. These were the marks of the kshatriya.