BREEDING SEASONS Most birds are not simply reluctant to breed at other times – they are physically incapable of doing so. This is because their reproductive system shuts down and shrinks, an adaptation that helps flying birds to save weight. The main exception to this rule are nomadic desert species, such as Zebra Finches and Budgerigars.
- source; ‘The Illustrated Encyclopedia of Birds’ issued in London
While I think the grammatical subject is ‘the main exception’, which needs the singular verb ‘is’ in the bold part above, why did the author use ‘are’? If the ‘are’ is to be allowed in this context, I guess he/she might have considered one of the following possibilities;
(1) the inversion applied; with the subject - ‘nomadic desert species’ and the complement - ‘the main exception’. (2) the notional agreement applied; some people consider the real subject to be ‘nomadic desert species’ semantically, while the formal subject is ‘the main exception’ (3) the principle of proximity applied; the verb ‘are’ (not 'is') has been finally agreed by the close plural complement ‘nomadic desert species’.
Even if one of the possibilities above has been applied, could it be grammatically accepted here?
Would appreciate your responses.