diff --git a/changes/bug22460_case1 b/changes/bug22460_case1
new file mode 100644
index 0000000000..cfe78ad791
--- /dev/null
+++ b/changes/bug22460_case1
@@ -0,0 +1,16 @@
+  o Major bugfixes (relays, key management):
+    - Regenerate link and authentication certificates whenever the key that
+      signs them changes; also, regenerate link certificates whenever the
+      signed key changes. Previously, these processes were only weakly
+      coupled, and we relays could (for minutes to hours) wind up with an
+      inconsistent set of keys and certificates, which other relays
+      would not accept. Fixes two cases of bug 22460; bugfix on
+      0.3.0.1-alpha.
+    - When sending an Ed25519 signing->link certificate in a CERTS cell,
+      send the certificate that matches the x509 certificate that we used
+      on the TLS connection. Previously, there was a race condition if
+      the TLS context rotated after we began the TLS handshake but
+      before we sent the CERTS cell. Fixes a case of bug 22460; bugfix
+      on 0.3.0.1-alpha.
+
+
diff --git a/src/or/connection_or.c b/src/or/connection_or.c
index b9ac9b2c5d..1e2d731f53 100644
--- a/src/or/connection_or.c
+++ b/src/or/connection_or.c
@@ -1855,6 +1855,9 @@ connection_init_or_handshake_state(or_connection_t *conn, int started_here)
   s->started_here = started_here ? 1 : 0;
   s->digest_sent_data = 1;
   s->digest_received_data = 1;
+  if (! started_here && get_current_link_cert_cert()) {
+    s->own_link_cert = tor_cert_dup(get_current_link_cert_cert());
+  }
   s->certs = or_handshake_certs_new();
   s->certs->started_here = s->started_here;
   return 0;
@@ -1869,6 +1872,7 @@ or_handshake_state_free(or_handshake_state_t *state)
   crypto_digest_free(state->digest_sent);
   crypto_digest_free(state->digest_received);
   or_handshake_certs_free(state->certs);
+  tor_cert_free(state->own_link_cert);
   memwipe(state, 0xBE, sizeof(or_handshake_state_t));
   tor_free(state);
 }
@@ -2234,7 +2238,8 @@ add_certs_cell_cert_helper(certs_cell_t *certs_cell,
 
 /** Add an encoded X509 cert (stored as <b>cert_len</b> bytes at
  * <b>cert_encoded</b>) to the trunnel certs_cell_t object that we are
- * building in <b>certs_cell</b>.  Set its type field to <b>cert_type</b>. */
+ * building in <b>certs_cell</b>.  Set its type field to <b>cert_type</b>.
+ * (If <b>cert</b> is NULL, take no action.) */
 static void
 add_x509_cert(certs_cell_t *certs_cell,
               uint8_t cert_type,
@@ -2252,7 +2257,7 @@ add_x509_cert(certs_cell_t *certs_cell,
 
 /** Add an Ed25519 cert from <b>cert</b> to the trunnel certs_cell_t object
  * that we are building in <b>certs_cell</b>.  Set its type field to
- * <b>cert_type</b>. */
+ * <b>cert_type</b>. (If <b>cert</b> is NULL, take no action.) */
 static void
 add_ed25519_cert(certs_cell_t *certs_cell,
                  uint8_t cert_type,
@@ -2315,9 +2320,10 @@ connection_or_send_certs_cell(or_connection_t *conn)
                    CERTTYPE_ED_ID_SIGN,
                    get_master_signing_key_cert());
   if (conn_in_server_mode) {
+    tor_assert_nonfatal(conn->handshake_state->own_link_cert);
     add_ed25519_cert(certs_cell,
                      CERTTYPE_ED_SIGN_LINK,
-                     get_current_link_cert_cert());
+                     conn->handshake_state->own_link_cert);
   } else {
     add_ed25519_cert(certs_cell,
                      CERTTYPE_ED_SIGN_AUTH,
diff --git a/src/or/main.c b/src/or/main.c
index bc7b3db2b9..3139381f30 100644
--- a/src/or/main.c
+++ b/src/or/main.c
@@ -1506,8 +1506,9 @@ check_ed_keys_callback(time_t now, const or_options_t *options)
 {
   if (server_mode(options)) {
     if (should_make_new_ed_keys(options, now)) {
-      if (load_ed_keys(options, now) < 0 ||
-          generate_ed_link_cert(options, now)) {
+      int new_signing_key = load_ed_keys(options, now);
+      if (new_signing_key < 0 ||
+          generate_ed_link_cert(options, now, new_signing_key > 0)) {
         log_err(LD_OR, "Unable to update Ed25519 keys!  Exiting.");
         tor_cleanup();
         exit(0);
@@ -1559,6 +1560,11 @@ rotate_x509_certificate_callback(time_t now, const or_options_t *options)
     log_err(LD_BUG, "Error reinitializing TLS context");
     tor_assert_unreached();
   }
+  if (generate_ed_link_cert(options, now, 1)) {
+    log_err(LD_OR, "Unable to update Ed25519->TLS link certificate for "
+            "new TLS context.");
+    tor_assert_unreached();
+  }
 
   /* We also make sure to rotate the TLS connections themselves if they've
    * been up for too long -- but that's done via is_bad_for_new_circs in
@@ -2298,8 +2304,9 @@ do_hup(void)
     /* Maybe we've been given a new ed25519 key or certificate?
      */
     time_t now = approx_time();
-    if (load_ed_keys(options, now) < 0 ||
-         generate_ed_link_cert(options, now)) {
+    int new_signing_key = load_ed_keys(options, now);
+    if (new_signing_key < 0 ||
+        generate_ed_link_cert(options, now, new_signing_key > 0)) {
       log_warn(LD_OR, "Problem reloading Ed25519 keys; still using old keys.");
     }
 
@@ -3627,7 +3634,7 @@ tor_main(int argc, char *argv[])
     result = do_main_loop();
     break;
   case CMD_KEYGEN:
-    result = load_ed_keys(get_options(), time(NULL));
+    result = load_ed_keys(get_options(), time(NULL)) < 0;
     break;
   case CMD_LIST_FINGERPRINT:
     result = do_list_fingerprint();
diff --git a/src/or/or.h b/src/or/or.h
index 0db9f23604..50e6e3e71b 100644
--- a/src/or/or.h
+++ b/src/or/or.h
@@ -1449,6 +1449,12 @@ typedef struct or_handshake_state_t {
   /* True iff we have sent a netinfo cell */
   unsigned int sent_netinfo : 1;
 
+  /** The signing->ed25519 link certificate corresponding to the x509
+   * certificate we used on the TLS connection (if this is a server-side
+   * connection). We make a copy of this here to prevent a race condition
+   * caused by TLS context rotation. */
+  struct tor_cert_st *own_link_cert;
+
   /** True iff we should feed outgoing cells into digest_sent and
    * digest_received respectively.
    *
diff --git a/src/or/router.c b/src/or/router.c
index e4fa72a283..f6b03cde27 100644
--- a/src/or/router.c
+++ b/src/or/router.c
@@ -906,7 +906,8 @@ init_keys(void)
   }
 
   /* 1d. Load all ed25519 keys */
-  if (load_ed_keys(options,now) < 0)
+  const int new_signing_key = load_ed_keys(options,now);
+  if (new_signing_key < 0)
     return -1;
 
   /* 2. Read onion key.  Make it if none is found. */
@@ -976,7 +977,7 @@ init_keys(void)
 
   /* 3b. Get an ed25519 link certificate.  Note that we need to do this
    * after we set up the TLS context */
-  if (generate_ed_link_cert(options, now) < 0) {
+  if (generate_ed_link_cert(options, now, new_signing_key > 0) < 0) {
     log_err(LD_GENERAL,"Couldn't make link cert");
     return -1;
   }
diff --git a/src/or/routerkeys.c b/src/or/routerkeys.c
index 611ac91397..1eb44db855 100644
--- a/src/or/routerkeys.c
+++ b/src/or/routerkeys.c
@@ -673,6 +673,9 @@ static time_t rsa_ed_crosscert_expiration = 0;
 /**
  * Running as a server: load, reload, or refresh our ed25519 keys and
  * certificates, creating and saving new ones as needed.
+ *
+ * Return -1 on failure; 0 on success if the signing key was not replaced;
+ * and 1 on success if the signing key was replaced.
  */
 int
 load_ed_keys(const or_options_t *options, time_t now)
@@ -685,6 +688,7 @@ load_ed_keys(const or_options_t *options, time_t now)
   const tor_cert_t *check_signing_cert = NULL;
   tor_cert_t *sign_cert = NULL;
   tor_cert_t *auth_cert = NULL;
+  int signing_key_changed = 0;
 
 #define FAIL(msg) do {                          \
     log_warn(LD_OR, (msg));                     \
@@ -722,7 +726,23 @@ load_ed_keys(const or_options_t *options, time_t now)
     use_signing = sign;
   }
 
+  if (use_signing) {
+    /* We loaded a signing key with its certificate.  */
+    if (! master_signing_key) {
+      /* We didn't know one before! */
+      signing_key_changed = 1;
+    } else if (! ed25519_pubkey_eq(&use_signing->pubkey,
+                                   &master_signing_key->pubkey) ||
+               ! tor_memeq(use_signing->seckey.seckey,
+                           master_signing_key->seckey.seckey,
+                           ED25519_SECKEY_LEN)) {
+      /* We loaded a different signing key than the one we knew before. */
+      signing_key_changed = 1;
+    }
+  }
+
   if (!use_signing && master_signing_key) {
+    /* We couldn't load a signing key, but we already had one loaded */
     check_signing_cert = signing_key_cert;
     use_signing = master_signing_key;
   }
@@ -882,6 +902,7 @@ load_ed_keys(const or_options_t *options, time_t now)
     if (!sign)
       FAIL("Missing signing key");
     use_signing = sign;
+    signing_key_changed = 1;
 
     tor_assert(sign_cert->signing_key_included);
     tor_assert(ed25519_pubkey_eq(&sign_cert->signing_key, &id->pubkey));
@@ -918,6 +939,7 @@ load_ed_keys(const or_options_t *options, time_t now)
   }
 
   if (!current_auth_key ||
+      signing_key_changed ||
       EXPIRES_SOON(auth_key_cert, options->TestingAuthKeySlop)) {
     auth = ed_key_new(use_signing, INIT_ED_KEY_NEEDCERT,
                       now,
@@ -945,7 +967,7 @@ load_ed_keys(const or_options_t *options, time_t now)
     SET_CERT(auth_key_cert, auth_cert);
   }
 
-  return 0;
+  return signing_key_changed;
  err:
   ed25519_keypair_free(id);
   ed25519_keypair_free(sign);
@@ -959,16 +981,18 @@ load_ed_keys(const or_options_t *options, time_t now)
  * Retrieve our currently-in-use Ed25519 link certificate and id certificate,
  * and, if they would expire soon (based on the time <b>now</b>, generate new
  * certificates (without embedding the public part of the signing key inside).
+ * If <b>force</b> is true, always generate a new certificate.
  *
- * The signed_key from the expiring certificate will be used to sign the new
- * key within newly generated X509 certificate.
+ * The signed_key from the current id->signing certificate will be used to
+ * sign the new key within newly generated X509 certificate.
  *
  * Returns -1 upon error.  Otherwise, returns 0 upon success (either when the
  * current certificate is still valid, or when a new certificate was
  * successfully generated).
  */
 int
-generate_ed_link_cert(const or_options_t *options, time_t now)
+generate_ed_link_cert(const or_options_t *options, time_t now,
+                      int force)
 {
   const tor_x509_cert_t *link_ = NULL, *id = NULL;
   tor_cert_t *link_cert = NULL;
@@ -980,7 +1004,8 @@ generate_ed_link_cert(const or_options_t *options, time_t now)
 
   const common_digests_t *digests = tor_x509_cert_get_cert_digests(link_);
 
-  if (link_cert_cert &&
+  if (force == 0 &&
+      link_cert_cert &&
       ! EXPIRES_SOON(link_cert_cert, options->TestingLinkKeySlop) &&
       fast_memeq(digests->d[DIGEST_SHA256], link_cert_cert->signed_key.pubkey,
                  DIGEST256_LEN)) {
@@ -1082,7 +1107,7 @@ init_mock_ed_keys(const crypto_pk_t *rsa_identity_key)
   MAKECERT(auth_key_cert,
            master_signing_key, current_auth_key, CERT_TYPE_SIGNING_AUTH, 0);
 
-  if (generate_ed_link_cert(get_options(), time(NULL)) < 0) {
+  if (generate_ed_link_cert(get_options(), time(NULL), 0) < 0) {
     log_warn(LD_BUG, "Couldn't make link certificate");
     goto err;
   }
diff --git a/src/or/routerkeys.h b/src/or/routerkeys.h
index d2027f4bbe..845abb4c70 100644
--- a/src/or/routerkeys.h
+++ b/src/or/routerkeys.h
@@ -66,7 +66,7 @@ MOCK_DECL(int, check_tap_onion_key_crosscert,(const uint8_t *crosscert,
 int load_ed_keys(const or_options_t *options, time_t now);
 int should_make_new_ed_keys(const or_options_t *options, const time_t now);
 
-int generate_ed_link_cert(const or_options_t *options, time_t now);
+int generate_ed_link_cert(const or_options_t *options, time_t now, int force);
 
 int read_encrypted_secret_key(ed25519_secret_key_t *out,
                               const char *fname);
diff --git a/src/test/test_link_handshake.c b/src/test/test_link_handshake.c
index b3e854ef76..ade7cf6998 100644
--- a/src/test/test_link_handshake.c
+++ b/src/test/test_link_handshake.c
@@ -911,6 +911,11 @@ test_link_handshake_send_authchallenge(void *arg)
   or_connection_t *c1 = or_connection_new(CONN_TYPE_OR, AF_INET);
   var_cell_t *cell1=NULL, *cell2=NULL;
 
+  crypto_pk_t *rsa0 = pk_generate(0), *rsa1 = pk_generate(1);
+  tt_int_op(tor_tls_context_init(TOR_TLS_CTX_IS_PUBLIC_SERVER,
+                                 rsa0, rsa1, 86400), ==, 0);
+  init_mock_ed_keys(rsa0);
+
   MOCK(connection_or_write_var_cell_to_buf, mock_write_var_cell);
 
   tt_int_op(connection_init_or_handshake_state(c1, 0), ==, 0);
@@ -936,6 +941,8 @@ test_link_handshake_send_authchallenge(void *arg)
   connection_free_(TO_CONN(c1));
   tor_free(cell1);
   tor_free(cell2);
+  crypto_pk_free(rsa0);
+  crypto_pk_free(rsa1);
 }
 
 typedef struct authchallenge_data_s {
diff --git a/src/test/test_routerkeys.c b/src/test/test_routerkeys.c
index 64692d28a0..13059267ac 100644
--- a/src/test/test_routerkeys.c
+++ b/src/test/test_routerkeys.c
@@ -450,8 +450,8 @@ test_routerkeys_ed_keys_init_all(void *arg)
 
   options->DataDirectory = dir;
 
-  tt_int_op(0, ==, load_ed_keys(options, now));
-  tt_int_op(0, ==, generate_ed_link_cert(options, now));
+  tt_int_op(1, ==, load_ed_keys(options, now));
+  tt_int_op(0, ==, generate_ed_link_cert(options, now, 0));
   tt_assert(get_master_identity_key());
   tt_assert(get_master_identity_key());
   tt_assert(get_master_signing_keypair());
@@ -466,7 +466,7 @@ test_routerkeys_ed_keys_init_all(void *arg)
 
   /* Call load_ed_keys again, but nothing has changed. */
   tt_int_op(0, ==, load_ed_keys(options, now));
-  tt_int_op(0, ==, generate_ed_link_cert(options, now));
+  tt_int_op(0, ==, generate_ed_link_cert(options, now, 0));
   tt_mem_op(&id, ==, get_master_identity_key(), sizeof(id));
   tt_mem_op(&sign, ==, get_master_signing_keypair(), sizeof(sign));
   tt_mem_op(&auth, ==, get_current_auth_keypair(), sizeof(auth));
@@ -474,8 +474,8 @@ test_routerkeys_ed_keys_init_all(void *arg)
 
   /* Force a reload: we make new link/auth keys. */
   routerkeys_free_all();
-  tt_int_op(0, ==, load_ed_keys(options, now));
-  tt_int_op(0, ==, generate_ed_link_cert(options, now));
+  tt_int_op(1, ==, load_ed_keys(options, now));
+  tt_int_op(0, ==, generate_ed_link_cert(options, now, 0));
   tt_mem_op(&id, ==, get_master_identity_key(), sizeof(id));
   tt_mem_op(&sign, ==, get_master_signing_keypair(), sizeof(sign));
   tt_assert(tor_cert_eq(link_cert, get_current_link_cert_cert()));
@@ -489,7 +489,7 @@ test_routerkeys_ed_keys_init_all(void *arg)
 
   /* Force a link/auth-key regeneration by advancing time. */
   tt_int_op(0, ==, load_ed_keys(options, now+3*86400));
-  tt_int_op(0, ==, generate_ed_link_cert(options, now+3*86400));
+  tt_int_op(0, ==, generate_ed_link_cert(options, now+3*86400, 0));
   tt_mem_op(&id, ==, get_master_identity_key(), sizeof(id));
   tt_mem_op(&sign, ==, get_master_signing_keypair(), sizeof(sign));
   tt_assert(! tor_cert_eq(link_cert, get_current_link_cert_cert()));
@@ -502,8 +502,8 @@ test_routerkeys_ed_keys_init_all(void *arg)
   memcpy(&auth, get_current_auth_keypair(), sizeof(auth));
 
   /* Force a signing-key regeneration by advancing time. */
-  tt_int_op(0, ==, load_ed_keys(options, now+100*86400));
-  tt_int_op(0, ==, generate_ed_link_cert(options, now+100*86400));
+  tt_int_op(1, ==, load_ed_keys(options, now+100*86400));
+  tt_int_op(0, ==, generate_ed_link_cert(options, now+100*86400, 0));
   tt_mem_op(&id, ==, get_master_identity_key(), sizeof(id));
   tt_mem_op(&sign, !=, get_master_signing_keypair(), sizeof(sign));
   tt_assert(! tor_cert_eq(link_cert, get_current_link_cert_cert()));
@@ -520,8 +520,8 @@ test_routerkeys_ed_keys_init_all(void *arg)
   routerkeys_free_all();
   unlink(get_fname("test_ed_keys_init_all/keys/"
                    "ed25519_master_id_secret_key"));
-  tt_int_op(0, ==, load_ed_keys(options, now));
-  tt_int_op(0, ==, generate_ed_link_cert(options, now));
+  tt_int_op(1, ==, load_ed_keys(options, now));
+  tt_int_op(0, ==, generate_ed_link_cert(options, now, 0));
   tt_mem_op(&id, ==, get_master_identity_key(), sizeof(id));
   tt_mem_op(&sign, ==, get_master_signing_keypair(), sizeof(sign));
   tt_assert(! tor_cert_eq(link_cert, get_current_link_cert_cert()));
diff --git a/src/test/test_shared_random.c b/src/test/test_shared_random.c
index d511f163e3..026a0f3825 100644
--- a/src/test/test_shared_random.c
+++ b/src/test/test_shared_random.c
@@ -48,7 +48,7 @@ init_authority_state(void)
   mock_cert = authority_cert_parse_from_string(AUTHORITY_CERT_1, NULL);
   tt_assert(mock_cert);
   options->AuthoritativeDir = 1;
-  tt_int_op(0, ==, load_ed_keys(options, time(NULL)));
+  tt_int_op(load_ed_keys(options, time(NULL)), OP_GE, 0);
   sr_state_init(0, 0);
   /* It's possible a commit has been generated in our state depending on
    * the phase we are currently in which uses "now" as the starting
@@ -286,7 +286,7 @@ test_sr_commit(void *arg)
     tt_assert(auth_cert);
 
     options->AuthoritativeDir = 1;
-    tt_int_op(0, ==, load_ed_keys(options, now));
+    tt_int_op(load_ed_keys(options, time(NULL)), OP_GE, 0);
   }
 
   /* Generate our commit object and validate it has the appropriate field