Firstly this is a very good question.
e.g. The authorization header or content type header. Which is absolutely required by the server to understand the request. This can differ from server to server.
This is less severe than 400. The request has reached the server. The server has acknowledged the request has got the basic structure right. But the information in the request body can't be parsed or understood.
e.g. Content-Type: application/xml
when request body is JSON.
Here's an article listing status codes and its use in REST APIs. https://metamug.com/article/status-codes-for-rest-api.php