Crustaceans and arachnids have respiratory pigments. Arthropods therefore replace their exoskeletons by undergoing ecdysis (moulting), or shedding the old exoskeleton after growing a new one that is not yet hardened. A few such species rely on females to find spermatophores that have already been deposited on the ground, but in most cases males only deposit spermatophores when complex courtship rituals look likely to be successful.  The mite Varroa destructor has become the largest single problem faced by beekeepers worldwide. However, most male terrestrial arthropods produce spermatophores, waterproof packets of sperm, which the females take into their bodies. Hemocyanin is the major respiratory pigment present in crustaceans, but hemoglobin is found in a few species and both are dissolved in the hemolymph rather than carried in cells. There had been competing proposals that arthropods were closely related to other groups such as nematodes, priapulids and tardigrades, but these remained minority views because it was difficult to specify in detail the relationships between these groups. , Working out the evolutionary stages by which all these different combinations could have appeared is so difficult that it has long been known as "the arthropod head problem". The hemolymph of onychophorans contains the respiratory pigment hemocyanins, as do arthropods. Hemocyanin is a respiratory pigment found in mollusks and arthropods.  Around the same time the aquatic, scorpion-like eurypterids became the largest ever arthropods, some as long as 2.5 metres (8.2 ft).  For example, Graham Budd's analyses of Kerygmachela in 1993 and of Opabinia in 1996 convinced him that these animals were similar to onychophorans and to various Early Cambrian "lobopods", and he presented an "evolutionary family tree" that showed these as "aunts" and "cousins" of all arthropods. Higher up the "family tree", the Annelida have traditionally been considered the closest relatives of the Panarthropoda, since both groups have segmented bodies, and the combination of these groups was labelled Articulata. Heart: The heart is composed of muscular ventricle and thin-walled auricle.  The cuticle of many crustaceans, beetle mites, and millipedes (except for bristly millipedes) is also biomineralized with calcium carbonate. The brain is in the head, encircling and mainly above the esophagus. It was assumed it was a non-discriminatory sediment feeder, processing whatever sediment came its way for food, but fossil findings hints that the last common ancestor of both arthropods and priapulida shared the same specialized mouth apparatus; a circular mouth with rings of teeth used for capturing prey and was therefore carnivorous. Head, appendages and respiratory organs in some cases, Circular and longitudinal muscles in both body wall and gut wall. Hemocyanins are copper-containing respiratory pigments which are found in many mollusks and arthropods (many crustaceans, some arachnids, and the horseshoe crab.Hemocyanins are colorless in the reduced, or deoxygenated, state and blue when exposed to air or to oxygen dissolved in the blood. , A contrary view was presented in 2003, when Jan Bergström and Xian-Guang Hou argued that, if arthropods were a "sister-group" to any of the anomalocarids, they must have lost and then re-evolved features that were well-developed in the anomalocarids. , Arthropods are invertebrates with segmented bodies and jointed limbs. This pigment is green and becomes red in the presence of oxygen.  Various groups of terrestrial arthropods have independently developed a different system: the end-product of nitrogen metabolism is uric acid, which can be excreted as dry material; the Malpighian tubule system filters the uric acid and other nitrogenous waste out of the blood in the hemocoel, and dumps these materials into the hindgut, from which they are expelled as feces. These would later fuse into a single pair of biramous appendages, with the upper branch acting as a gill while the lower branch was used for locomotion. , A few arthropods, such as barnacles, are hermaphroditic, that is, each can have the organs of both sexes. Small species often do not have any, since their high ratio of surface area to volume enables simple diffusion through the body surface to supply enough oxygen. At this point, the new one is wrinkled and so soft that the animal cannot support itself and finds it very difficult to move, and the new endocuticle has not yet formed. , Living arthropods have paired main nerve cords running along their bodies below the gut, and in each segment the cords form a pair of ganglia from which sensory and motor nerves run to other parts of the segment. In crustaceans and in arachnids hemocyanin is the respiratory pigment. Along the heart run a series of paired ostia, non-return valves that allow blood to enter the heart but prevent it from leaving before it reaches the front. , Although arthropods are the most numerous phylum on Earth, and thousands of arthropod species are venomous, they inflict relatively few serious bites and stings on humans. (1979). If the Ecdysozoa hypothesis is correct, then segmentation of arthropods and annelids either has evolved convergently or has been inherited from a much older ancestor and subsequently lost in several other lineages, such as the non-arthropod members of the Ecdysozoa. Moulting may be responsible for 80 to 90% of all arthropod deaths.  Compound eyes have a wide field of view, and can detect fast movement and, in some cases, the polarization of light. The respiratory pigment is haemocyanin, which contains both iron and copper.