Onychophorans are carnivorous, terrestrial invertebrates that occur in tropical and temperate forests of the Southern Hemisphere and around the Equator. Together with tardigrades, onychophorans are regarded as one of the closest relatives of arthropods. One of the most peculiar features of onychophorans is their hunting and feeding behavior. These animals secrete a sticky slime, which is ejected via a pair of slime-papillae, to entangle the prey. After the prey has been immobilized, its cuticle is punctured using a pair of jaws located within the mouth. These jaws constitute internalized appendages of the second body segment and are innervated by the deutocerebrum; thus, they are homologous to the chelicerae of chelicerates, and to the (first) antennae of myriapods, crustaceans, and insects. The jaws are also serial homologs of the paired claws associated with each walking limb of the trunk. The structure of the jaws is similar in representatives of the two major onychophoran subgroups, the Peripatidae and Peripatopsidae. Each jaw is characterized by an outer and an inner blade; while the outer blade consists only of a large principal tooth and up to three accessory teeth, the inner blade bears numerous additional denticles. These denticles are separated from the remaining part of the inner jaw by a diastema and a soft membrane only in peripatids. The onychophoran jaws are associated with large apodemes and specialized muscles that enable their movement. In contrast to the mandibles of arthropods, the onychophoran jaws are moved along, rather than perpendicular to, the main axis of the body. Our elemental analysis reveals an increased incorporation of calcium at the tip of each blade, which might provide rigidity, whereas there is no evidence for incorporation of metal or prominent mineralization. Stability of the jaw might be further facilitated by the cone-in-cone organization of its cuticle, as each blade consists of several stacked, cuticular elements. In this work, we summarize current knowledge on the jaws of onychophorans, which are a characteristic feature of these animals.
Arthropods are well known to biomineralize and metallize their exoskeletons with naturally occurring elements that function to stabilize the protein component and add hardness to surfaces that endure wear. In this study, we provide the first description of the trace elements in the cuticle of a marine intertidal pseudoscorpion, Halobisium occidentale, using energy‐dispersive x‐ray spectroscopy. We characterized the trace element composition of six regions on four specimens: the prosoma, opisthosoma, tarsal claws, arolia, chelicerae, and pedipalps. In addition to the elements C, O, and N that make up the α‐chitin component of the cuticle, we found 11 trace elements across the body, but only five of these elements were present at significant levels (≥1% wt): Ni, P, Al, Zn, and Fe. The only trace element on both tagmata is P, while the appendages and their structures contain varying amounts of other elements. The tarsal claws are supplemented with Zn and Ni, while the adhesive arolia contain either Ni or P. The pedipalps are enriched with Al along their proximodistal axis, with P, Zn, and Fe present only around the venom pore. The chelicerae have P, Zn, and Fe present only on the distal regions. This study confirms that pseudoscorpions, like many other arthropods, enrich their cuticle with specific elements at precise locations that are important in predation (cheliceral fingers, pedipalps, venom pore) and locomotion (tarsal claws, arolia). This is also the first study to reveal the presence of Al and Ni in any significant quantities in the arthropod exoskeleton.
The arthropod cuticle is a structurally diverse secretion that is largely composed of lipids, proteins, and α‐chitin that function together in protection, prey capture, and as a skeletal framework for efficient and diverse means of locomotion. Aquatic, aerial, and terrestrial arthropods are well known to enrich their cuticles with trace elements that are proposed to strengthen or otherwise enhance specific regions of the exoskeleton. In this study, we provide evidence that whipscorpions (vinegaroons) enrich their exoskeleton with up to 14 trace elements. We use energy‐dispersive x‐ray spectroscopy (EDS) to provide the first comparisons of trace element composition in two species of Thelyphonida: Mastigoproctus giganteus (female) and Typopeltis dalyi (female and male). We analyzed the chemical composition of eight regions that have sensory, locomotory, taxonomic, protective, and/or predatory significance: the flagellum, prosoma, tarsal claws of the walking legs, tarsi of the antenniform legs, chelicerae, and the terminal three segments of the pedipalps. Our results reveal the presence of 14 trace elements across both species (12 elements in T. dalyi, 10 elements in M. giganteus), and only four elements are present at significant levels (≥1% weight): Si, Cl, Ca, and Zn. The flagellum, antenniform leg tarsi, and prosoma keel lack these elements in both species, while the chelicerae of all species are enriched with Ca, Zn, and Cl, and the tarsal claws are enriched with Zn. Significantly, we note the presence of Si in the prosomal carapace (but not the keel) of males and females of Typopeltis only, which appears to be the first evidence of this transition metal in the arthropod exoskeleton. We discuss the significance of these chemical enrichments in whipscorpions and provide hypotheses about their functional significance.
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