Oligomeric larvae of the pycnogonids revisited

Organization and ultrastructure of the protonymphon larva were never adequately described, despite it being the common larval type of the enigmatic sea spiders and the only example of oligosegmented life stage among recent chelicerates. We have made a comprehensive examination of the newly hatched free-living protonymphons of Nymphon brevirostre using SEM, TEM, light, and confocal microscopy. Although fairly typical in their broad characters, protonymphon larvae have a number of unique and unexpected traits. Body cavity, already present at this stage, is lined with extracellular matrix and thus is conclusively identified as primary body cavity. Central nervous system includes four postocular neuromeres arranged in three ganglia: supraesophageal, subesophageal, and the first ganglion of the ventral nerve cord. Examination of the sensory organs revealed unusually organized eyes, mechanoreceptors, and chemoreceptors. We have uncovered a mixed sensory-secretory nature of chelar glands and proposed possible modalities of its receptory part. We gave first descriptions of the complex ultrastructure of three secretory organs (spinning glands, slit-like organs, proboscis glands) and hypothesized on their mode of functioning. Comparisons with another oligomeric larva, for example, nauplius, revealed discrepancies in the segmentation of these animals. Although both larvae are externally unsegmented and bear three pairs of homologous appendages, the protonymphon body includes a fourth segment of the prospective walking legs which is absent in nauplius.