Tiny but complex - interactive 3D visualization of the interstitial acochlidian gastropod Pseudunela cornuta (Challis, 1970)

Background: Mesopsammic acochlidians are small, and organ complexity may be strongly reduced (regressive evolution by progenesis), especially in microhedylacean species. The marine interstitial hedylopsacean Pseudunela cornuta (Challis, 1970), however, was suggested as having a complex reproductive system resembling that of much larger, limnic and benthic species. The present study aims to reconstruct the detailed anatomy and true complexity of P. cornuta from serial, semithin histological sections by using modern computer-based 3D visualization with Amira software, and to explain it in an evolutionary context. Results: Our results demonstrate considerable discordance with the original species description, which was based solely on paraffin sections. Here, we show that the nervous system of P. cornuta has paired rhinophoral, optic and gastro-oesophageal ganglia, three distinct ganglia on the visceral nerve cord, and a putative osphradial ganglion, while anterior accessory ganglia are absent. The presence of an anal genital cloaca is clearly rejected and the anus, nephropore and gonopore open separately to the exterior; the circulatory and excretory systems are well-differentiated, including a two-chambered heart and a complex kidney with a long, looped nephroduct; the special androdiaulic reproductive system shows two allosperm receptacles, three nidamental glands, a cavity with unknown function, as well as highly complex anterior copulatory organs with two separate glandular and impregnatory systems including a penial stylet that measures approximately a third of the whole length of the preserved specimen. Conclusion: In spite of its small body size, the interstitial hermaphroditic P. cornuta shows high complexity regarding all major organ systems; the excretory system is as differentiated as in species of the sister clade, the limnic and much larger Acochlidiidae, and the reproductive system is by far the most elaborated one ever observed in a mesopsammic gastropod, though functionally not yet fully understood. Such organ complexity as shown herein by interactive 3D visualization is not plesiomorphically maintained from a larger, benthic ancestor, but newly evolved within small marine hedylopsacean ancestors of P. cornuta. The common picture of general organ regression within mesopsammic acochlidians thus is valid for microhedylacean species only