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What do the SMTP error / reply codes mean?

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Here is a reference table of SMTP error / reply codes and a brief description of their meaning:

Code Description
211 System status or system help reply.
214 Help message.
220 Domain service ready - ready to start TLS.
221 Domain service closing transmission channel.
250 OK - queuing for node started - requested mail action okay, completed.
251 OK - no messages waiting for node - user not local will forward to forward path.
252 OK - pending messages for node started - cannot VRFY user but will accept message and attempt delivery.
253 OK - messages pending - messages for node started.
354 Start mail input - end with <CRLF>.<CRLF>.
355 Octet-offset is the transaction offset.
421 Domain service not available - closing transmission channel.
432 A password transition is needed.
450 Requested mail action not taken: mailbox unavailable - ATRN request refused.
451 Requested action aborted - local error in processing - unable to process ATRN request.
452 Requested action not taken - insufficient system storage.
453 You have no mail.
454 TLS not available due to temporary reason - encryption is required for requested authentication mechanism.
458 Unable to queue messages for node
459 Node not allowed: reason.
500 Command not recognized 'command' - syntax error.
501 Syntax error - no parameters allowed.
502 Command not implemented.
503 Bad sequence of commands.
504 Command parameter not implemented.
521 Machine does not accept mail.
530 Must issue a STARTTLS command first - encryption required for requested authentication mechanism.
534 Authentication mechanism is too weak.
538 Encryption required for requested authentication mechanism.
550 Requested action not taken - mailbox unavailable.
551 User not local - please try forward path.
552 Requested mail action aborted - exceeded storage allocation.
553 Requested action not taken - mailbox name not allowed.
554 Transaction failed.

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