What does one call a comedian whose repertoire relies on sexual puns about feet? Answer: an ancient Athenian.

In the theaters of that cosmopolitan city in the fifth and fourth centuries BCE, Greek comic actors could always get a laugh with the popular pun that related feet to the male genitalia. For centuries, whether male or female, feet were associated with love and desire in ancient Greece, and evidence of their role abounds in literature and art.

"The foot left its imprint on Greek erotic thought from Homer to the rise of Christianity," said Daniel Levine, professor of classical studies at the University of Arkansas.

In "Eraton Bama ('Her Lovely Footstep'): The Erotics of Feet in Ancient Greece," Levine draws on archeological and literary sources to extend recent scholarship on the erotic aspects of feet and shoes to a consideration of ancient Greek culture. His work is a chapter in "Body Language in the Greek and Roman Worlds," edited by Douglas Cairns of University of Edinburgh.

The fondness of Athenian comedians for crude puns about feet reflects a rich tradition of associating feet with love, desire and fertility. In Greek epics, Levine observes, female beauty often resides in fine ankles and lovely feet. When the poet Hesiod describes the birth of Aphrodite, he stresses her shapely feet that give rise to new grass when she first steps on land, accompanied by Eros (the god of love) and Himeros (the god of desire).

"In addition to beautiful female ankles symbolizing the erotic ideal, the Greeks also linked men's feet to eros in poetry, arts and myth," Levine writes. "When men and women join in matrimony, feet and shoes play important ritual and artistic roles."

A preliminary ritual was the tying of the bride's sandals. As depicted on the decoration of an Attic Red-Figure vessel from the fifth century BCE, Eros stoops to bind a bride's sandal while a human attendant presents her bridal crown. The essayist Lucian describes a painting of Alexander's wedding in which several Eros figures assist the bride after the wedding ceremony. One smiling Eros lifts her veil while another removes her sandal to prepare her for the marriage bed.

Similarly, a fragment of Sappho's poetry plays with the association between feet and sexuality. Outside the wedding chamber, "The doorkeeper has feet seven fathoms long."

The morning after the consummation of the marriage, gifts were presented to the newlyweds, traditionally including footwear. The bride's gifts provided her with the tools to maintain her seductive, wedding-night beauty - jewelry, perfume, cosmetics and sandals.

For centuries, passages from Greek literature praised the beauty not just of feet, but of the mark they left on the earth. The title of Levine's chapter comes from Sappho's longing for her lover's lovely footstep. In poems and letters, lovers long to kiss footsteps or to step in their beloveds' bare footprints.

Levine writes that in the archaic and classical periods, "the relation between eros and the foot existed not only in the literary world of epic, the clever wordplay of the theater, the lyric poetry of the aristoi, and the sophisticated world of the symposium. Greeks from all walks of life seem to have seen the foot's erotic symbolism."

In a Christian text of the late second century, Clement of Alexandria "condemns the unseemly practice of erotic messages which some women create with inscribed sandals: 'And many women engrave on their sandals erotic greetings, so that from their walking they put a regular pattern on the ground, and they stamp out on it the whorish nature of their intentions by their footsteps.'"

"Most of the time a foot is just a foot," Levine notes. However, the impact of the Greek appreciation for the erotic aspect of feet, ankles, sandals and footprints had a broad and diverse effect on their culture and world view. The verb "to walk" and the word for 'plain,' which derives from the word for foot, both doubled as sexual slang. Incised graffiti in the ancient city of Thera include phrases related to sexual intercourse with foot-shaped forms walking over them, a symbolic representation of the earth as "a place where men sexually 'tread'."

In his introduction to "Body Language in the Greek and Roman Worlds," Cairns notes that Levine has published "path-breaking studies of Homeric laughter" that are important contributions to the burgeoning interest by classical scholars in nonverbal communication in the ancient world. Levine's studies are cited in another chapter in the book by Michael Clarke of the National University of Ireland, Maynooth, on ancient Greek smiles.