Mechanisms of light organ occlusion in flashlight fishes, family Anomalopidae (Teleostei:Beryciformes),and the evolution of the group

The circumtropical, nocturnal, shore-fish family Anomalopidae is characterized by a subocular luminous organ containing symbiotic luminous bacteria. The five known species are placed in four genera, one of which is new. Phthanophaneron is restricted to the eastern Pacific, Kryptophanaron to the western Atlantic, Photoblepharon is Indo-West Pacific in distribution and Anomalops is west Pacific. The symbiotic bacteria emit light continuously, and two superficially different mechanisms of occluding the glowing face of the organ are found. In Photoblepharon a black shutter of elastic skin is drawn up over the face of the organ, whereas in Anomalops the organ is rotated downward, so that only the heavily pigmented back of the organ is exposed. In Phthanophaneron and Kryptophanaron, both rotational and shutter mechanisms are present. Elucidation of the structures and linkages involved in light-organ occlusion reveals that the superficially different mechanisms are based on a common functional complex. In all four genera, the light organ is supported by a cartilaginous cup that articulatrs anteriorly with a cartilaginous stalk. Motive power for both the shutter and rotational mechanisms is supplied by the adductor mandibulae through a complex biomechanical linkage involving the ethmomaxillary ligament and a ligament unique to anomalopids, the Ligament of Diogenes. The structures involved in shuttrr erection and organ rotation are illustrated and described in detail for Photoblepharon and Anomalops and are compared with those in the other two forms; a functional hypothesis is advanced. Extrafamilial relationships of the Anomalopidae are discussed, and a hypothesis of the phylogenetic relationships of the four genera is derived from a cladistic analysis involving 19 non-light-organ characters and corroborated by some light-organ characters. Most characters associated with the light-organ complex cannot be polarized by conventional outgroup comparison, and the evolution of the light organ occlusion mechanisms is interpreted in light of the hypothesized phylogeny and a hypothesized ancestral mechanism. We propose that the common ancestor of anomalopids possessed a forced rotational mechanism like that of Phthanophaneron and Kryptophanaron. This was refined to a more efficient flipping rotational mechanism in Anomalops, the sister group of the lineage comprising the other three genera, within which the shutter mechanism was progressively refined. The ostensibly unnecessary complexity of the shutter mechanism is apparently a result of functional-morphological constraints imposed on the system by the pre-existence of a rotational mechanism. A brief zoogeographic scenario is proposed.