The subphylum (or phylum) Chelicerata (pronounced /kəˌlɪsəˈreɪtə/ or /kəˌlɪsəˈrɑːtə/) constitutes one of the major subdivisions of the phylum (or superphylum) Arthropoda, and includes horseshoe crabs, scorpions, spiders and mites. They originated as marine animals, possibly in the Cambrian period, but the first confirmed chelicerate fossils, eurypterids, date from 445 million years ago in the Late Ordovician period. The surviving marine species include the four species of Xiphosurans (horseshoe crabs), and possibly the 1,300 species of Pycnogonida (sea spiders), if the latter are chelicerates. On the other hand, there are over 77,000 well-identified species of air-breathing chelicerates, and there may be about 500,000 unidentified species.
Like all arthropods, chelicerates have segmented bodies with jointed limbs, all covered in a cuticle made of chitin and proteins. The chelicerate bauplan consists of two tagmata, the cephalothorax and the abdomen, except that mites have lost a visible division between these sections. The chelicerae, which give the group its name, are the only appendages that appear before the mouth. In most sub-groups they are modest pincers used in feeding. However, spiders' chelicerae form fangs which in most species are used to inject venom into their prey. The group has the open circulatory system typical of the arthropods, in which a tube-like heart pumps blood through the hemocoel, which is the major body cavity. Marine chelicerates have gills, while the air-breathing forms generally have both book lungs and tracheae. In general the ganglia of living chelicerates' central nervous systems fuse into large masses in the cephalothorax, but there are wide variations and this fusion is very limited in the Mesothelae, which are regarded as the oldest and most primitive group of spiders. Most chelicerates rely on modified bristles for touch and for information about vibrations, air currents, and chemical changes in their environment. The most active hunting spiders also have very acute eyesight.
Chelicerates were originally predators, but the group has diversified to use all the major feeding strategies: predation, parasitism, herbivory, scavenging and eating decaying organic matter. Although harvestmen can digest solid food, the guts of most modern chelicerates are too narrow for this, and they generally liquidize their food by grinding it with their chelicerae and pedipalps and flooding it with digestive enzymes. To conserve water, air-breathing chelicerates excrete waste as solids that are removed from their blood by Malpighian tubules, structures which also evolved independently in insects. While the marine horseshoe crabs rely on external fertilization, air-breathing chelicerates use internal but usually indirect fertilization. Predatory species generally use elaborate courtship rituals to prevent males from being eaten before they can mate. Most lay eggs that hatch as what look like miniature adults, but all scorpions and a few species of mites keep the eggs inside their bodies until the young emerge. In most chelicerate species the young have to fend for themselves, but in scorpions and some species of spider the females protect and feed their young.
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