At high elevatiol), marsh marigold used more water per unit leaf area (40 lbs ft -2 ) than did willows or sedges (30 to 33 lbs ft -2 ), but its annual TWU was less than the other species because of its lesser total leaf area.
(its annual TWU; was; less than the other species; because of its lesser total leaf area)
its annual TWU
was
less than the other species because of its lesser total leaf area