In answer to your suggestion, we have a router with NAT configured on it between the ISP modem and the domain. Further investigation reveals there were two problems.
The first was that the severs except the DC1 all decided to start using IPV6 to the non-authoritative DNS as the primary means of resolving queries. I resolved that by turning off IPV6 on the domain servers, flushed the DNS resolver cache on all the servers, and deleted all the IPV6 cnames that had built up. They answer first properly now on all queries from the servers.
The second issue is that the workstations are now listing an IPV6 DNS server in the 1st position and the local domain servers in the 2nd and 3rd position. DHCP is turned off on the router, and no IPV6 address or scope is listed in the domain DHCP servers. I have no idea where it came, or is coming from. The DNS server IP that answers is registered to Network Solutions, which is where the mail server is for the domain, and where the company webpage is hosted.
The way it worked for years was the local computers queried the local controller 1. If it failed they queried the backup controller 2. If neither could resolve a name when asked they used a forwarder. I have no idea how an IVP6 DNS entry could put itself in first position when DHCP is configured otherwise.