During a parasitological survey of the cockscomb hydrobe, Littoridinops monroensis (Gastropoda: Cochliopidae), sympatric saltmarsh fishes and the American alligator, Alligator mississippiensis (Alligatoridae) within a saltmarsh pond in the Bon Secour Bay (Mobile Bay, Alabama, north-central Gulf of America) and other localities, we collected specimens of Proctocaecum diploporum (Stunkard, 1931) Baugh, 1957 and Proctocaecum coronarium (Cobbold, 1861) Brooks, 1980 (Cryptogonimidae). Fish scales were infected with 2 metacercarial cyst size classes, a small (<200 μm) and a large (>240 μm), suggesting the presence of distinct but closely related sympatric cryptogonimids. Morphological and nucleotide evidence demonstrated that P. diploporum (small cyst) and P. coronarium (large cyst), previously regarded as taxonomic synonyms, share the same first intermediate host (cockscomb hydrobe), are encysted within the scales of the same fish host species, and mature in the intestine of the same definitive host. The taxonomy of these trematodes and that of another similar American alligator trematode, Caimanicola pavida (Brooks and Overstreet, 1977) Brooks, 1980, needed reconsideration. Herein, we describe the first complete and naturally occurring life cycle of a reptile-infecting cryptogonimid, provide the first nucleotide data for a species of Proctocaecum, taxonomically reassess both P. diploporum and P. coronarium, and designate a neotype for P. coronarium. We consider C. pavida a junior subjective synonym of P. coronarium. Phylogenetically related and sympatric trematodes that use the same invertebrate and vertebrate hosts such as these Proctocaecum spp. could be a good model to further explore the mechanisms of speciation and host specificity among trematodes with complex life cycles.