Reassessing enigmatic mussel declines in the United States
Freshwater mussels have disappeared from many U.S. streams since the 1960s. These declines are enigmatic: there are no clear causes and other components of aquatic communities appear unaffected. I review the characteristics, spatial occurrence, timing, and potential causes of enigmatic mussel declines. They share some or all of the following characteristics: (1) fauna-wide collapse, affecting all species; (2) recruitment failure, leading to a senescent fauna; (3) no well-documented impact sufficient to affect all species rapidly; (4) specific to mussels; (5) recent occurrence, since the 1960s; (6) rapid action, often leading to faunal collapse within 10 yr; and (7) upstream progression in some cases. Enigmatic declines are largely restricted to upland regions south of maximum Pleistocene glaciation and north or west of the Gulf and Atlantic coastal plains, and they appear restricted to small- to medium-sized streams. In contrast, mussel declines with different characteristics are reported nationwide. Their consistent characteristics, restricted spatial occurrence, and similar timing suggest that enigmatic declines represent a distinct, diagnosable phenomenon. Many commonly invoked factors are not plausible explanations for enigmatic declines, and others are vague or poorly supported. Other factors are plausible in some cases (e.g., agricultural effects) but cannot explain declines across the affected area. I identified only two factors that could broadly explain enigmatic declines: disease and introduction of Corbicula fluminea, but these factors are poorly understood. The occurrence of enigmatic declines overlies the region with the highest mussel species richness on Earth, but I believe their severity and importance are underappreciated. Streams affected by enigmatic declines are vital research and management opportunities, deserving of increased attention; I propose ways that research can be focused to rigorously evaluate the specific mechanisms for these declines. Until we understand the causes of enigmatic declines, mussel conservation in affected areas is substantially hamstrung.