diff --git a/jose/jwt.py b/jose/jwt.py
index f47e4dd..308c564 100644
--- a/jose/jwt.py
+++ b/jose/jwt.py
@@ -363,7 +363,7 @@ def _validate_aud(claims, audience=None):
         raise JWTClaimsError("Invalid claim format in token")
     if any(not isinstance(c, str) for c in audience_claims):
         raise JWTClaimsError("Invalid claim format in token")
-    if audience not in audience_claims:
+    if audience is not None and audience not in audience_claims:
         raise JWTClaimsError("Invalid audience")
 
 
diff --git a/tests/test_jwt.py b/tests/test_jwt.py
index f9d54cd..6d81f95 100644
--- a/tests/test_jwt.py
+++ b/tests/test_jwt.py
@@ -359,6 +359,38 @@ def test_aud_empty_claim(self, claims, key):
         token = jwt.encode(claims, key)
         jwt.decode(token, key, audience=aud)
 
+    def test_aud_none_with_aud_claim_present(self, key):
+        # RFC 7519 4.1.3: a principal only has to identify itself in the
+        # "aud" claim when it verifies as an intended recipient. A caller
+        # that passes audience=None is not identifying itself, so a present
+        # "aud" claim must not cause verification to fail. See issue #389.
+        claims = {"aud": "audience"}
+
+        token = jwt.encode(claims, key)
+        jwt.decode(token, key, audience=None)
+
+    def test_aud_none_with_aud_list_claim_present(self, key):
+        claims = {"aud": ["audience", "another"]}
+
+        token = jwt.encode(claims, key)
+        jwt.decode(token, key, audience=None)
+
+    def test_aud_none_default_argument(self, key):
+        # audience defaults to None, so omitting it entirely must also pass
+        # when the token carries an "aud" claim.
+        claims = {"aud": "audience"}
+
+        token = jwt.encode(claims, key)
+        jwt.decode(token, key)
+
+    def test_aud_verify_disabled_with_aud_claim(self, key):
+        # verify_aud=False must still skip the check even when an audience
+        # is supplied and would otherwise mismatch.
+        claims = {"aud": "audience"}
+
+        token = jwt.encode(claims, key)
+        jwt.decode(token, key, audience="other", options={"verify_aud": False})
+
     def test_aud_not_string_or_list(self, key):
         aud = 1