Tarsal sensory structures in harvestmen (Arachnida, Opiliones): functional morphology, evolution and their use in systematics

Harvestmen (Arachnida, Opiliones) are especially dependent on contact chemoreception and are dependent on high humidity levels and amenable temperatures. However, knowledge on the sensory structures (sensilla) that detect such stimuli is limited in harvestmen when compared with other arachnid orders. Besides, there are few studies investigating the internal morphology of these structures -which is important for inferring function - and the potential of sensilla for systematics. To refine the knowledge on chemoreceptive sensilla and investigate the identity of hygro- and thermoreceptors, we investigated the tarsal sensilla of all leg pairs of the species Heteromitobates discolor (Laniatores, Gonyleptidae), using scanning electron microscopy (SEM) and transmission electron microscopy. To determine scope of the results obtained with H. discolor and to test the use of some structures for systematics, we surveyed (with SEM) species in all four suborders of Opiliones (Cyphophthalmi, Eupnoi, Dyspnoi and Laniatores), with focus in Laniatores (suborder with 2/3 of harvestmen species). In the first part, we provide the first morphological evidence of olfactory receptors in a species of Laniatores (H. discolor), showing that olfactory sensilla are abundant and diverse. Also, we show evidence (with SEM) of olfactory sensilla on 17 families of Laniatores, which suggests that olfaction is more important for Laniatores than previously considered. In the second part, we provide the first morphological evidence of hygro- and thermoreceptors in harvestmen (in H. discolor) discussing the functioning mechanisms of these structures and their morphological inter-association on the distal-most part of leg pairs I and II. Finally, we show that these putative hygro- thermoreceptive sensilla are widespread in species of Laniatores (28 families) and that comparable structures occur on species of the suborders Cyphophthalmi, Eupnoi and Dyspnoi. With an ancestral state reconstruction on a compiled phylogeny of Opiliones, we show that morphological variations on these sensilla are informative on several levels of phylogenetic relationships in Opiliones. This study constributes for the basic knowledge on the cellular anatomy of Opiliones sensilla, refining the function of tarsal sensillar types and providing a base for their use in systematics (AU)