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For print titles, generally capitalize all words (nouns, pronouns, verbs, adverbs, and adjectives) except conjunctions and prepositions of three letters or fewer, articles, and the to of infinitives unless aesthetically displeasing. Don't capitalize a, an, and, at, but, by, for, in, of, on, or, so, the, to, and up. Capitalize Too and YetPrepositions are capitalized when they are part of a verb, such as "believe in," "brought up," or "take off." 

Capitalize the first and last words of a title, even if they would not normally be capped. Words paralleling others that are capped should themselves be capped.

What Shape Is Your Population In?
What Timber Is Used For
To Be or Not To Be

Animals In and Around Ponds
Neither Plant Nor Animal
Use a colon or dash when one element amplifies another and a comma when one element is in apposition to another.

Chinatown, the Gilded Ghetto
Petra, Ancient City of Stone
Madagascar: A World Apart

Remnants: The Last Jews of Poland
Baltistan—The 20th Century Comes to Shangri-la
Senegambia—A Now and Future Nation
In hyphenated compounds, each word may be capped, depending on appearance.

Canada's Fur-Trading Empire
London's Eco-Friendly Pickle
but Along Afghanistan's War-torn Frontier

For digital articles the preferred style for articles and subheads is sentence case.