The complex trophic anatomy of silver carp, Hypophthalmichthys molitrix, highlighting a novel type of epibranchial organ

Hypophthalmichthys molitrix, silver carp, is an invasive Asian carp that has become increasingly widespread and ecologically destructive within the upper Mississippi River Basin. Its complex trophic anatomy may help explain the apparent efficiency with which they consume phytoplankton, outcompeting native filter feeders. This cypriniform species is characterized by trophic synapomorphies that include a palatal organ, loss of upper pharyngeal jaws, and a hypertrophied lower pharyngeal jaw. However, in silver carp these structures have become greatly modified and diverge from the more basal condition that characterizes species such as goldfish. The trophic apparatus of silver carp is composed of discrete structures that are functionally coupled: filtering plates, paired epibranchial organs (EBO), a modified palatal organ composed of large muscular folds that interdigitate with the filtering plates, and hypertrophied lower pharyngeal jaws and teeth. The filtering plates fill a significant portion of the buccal cavity, especially since the distal parts of these filtering plates make up a key component of the EBOs. EBOs, food aggregating structures found in many teleosts, are thought to have independently evolved at least six times. Ranging in complexity from small slits on the dorsal wall of the pharyngeal cavity to exceedingly intricate spiraling structures, EBOs are morphologically diverse among filter-feeding fishes. Despite this morphological diversity and broad taxonomic distribution, little is known regarding the functional anatomy of the EBO. Moreover, the EBO in silver carp is distinct from the organs previously described in other species, being created by four independent pharyngeal involutions (instead of the more typical one or two) that form spiral-shaped pharyngeal tubes surrounded by circumferential muscle. On each side of the head greatly hypertrophied hyomandibulae and opercles are connected to the anterior cartilaginous caps of the bilateral EBOs via enlarged muscles. Given that these fish are pump filter feeders we hypothesize that the opercula may compress and expand the EBOs during pumping causing food to be moved posteriorly toward the pharyngeal jaws.