section_path: Evolutionary history/Evolutionary relationships to other animal phyla
section_name: Evolutionary relationships to other animal phyla
paragraph_text: Evolutionary relationships to other animal phyla
  
  From 1952 to 1977, zoologist Sidnie Manton and others argued that arthropods are polyphyletic, in other words, that they do not share a common ancestor that was itself an arthropod. Instead, they proposed that three separate groups of "arthropods" evolved separately from common worm-like ancestors: the chelicerates, including spiders and scorpions; the crustaceans; and the uniramia, consisting of onychophorans, myriapods and hexapods. These arguments usually bypassed trilobites, as the evolutionary relationships of this class were unclear. Proponents of polyphyly argued the following: that the similarities between these groups are the results of convergent evolution, as natural consequences of having rigid, segmented exoskeletons; that the three groups use different chemical means of hardening the cuticle; that there were significant differences in the construction of their compound eyes; that it is hard to see how such different configurations of segments and appendages in the head could have evolved from the same ancestor; and that crustaceans have biramous limbs with separate gill and leg branches, while the other two groups have uniramous limbs in which the single branch serves as a leg.
  
  Further analysis and discoveries in the 1990s reversed this view, and led to acceptance that arthropods are monophyletic, in other words they are inferred to share a common ancestor that was itself an arthropod. 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. These changes made the scope of the term "arthropod" unclear, and Claus Nielsen proposed that the wider group should be labelled " Panarthropoda " ("all the arthropods") while the animals with jointed limbs and hardened cuticles should be called "Euarthropoda" ("true arthropods").
  
  A contrary view was presented in 2003, when Jan Bergström and Hou Xian-guang 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. The earliest known arthropods ate mud in order to extract food particles from it, and possessed variable numbers of segments with unspecialized appendages that functioned as both gills and legs. Anomalocarids were, by the standards of the time, huge and sophisticated predators with specialized mouths and grasping appendages, fixed numbers of segments some of which were specialized, tail fins, and gills that were very different from those of arthropods. In 2006, they suggested that arthropods were more closely related to lobopods and tardigrades than to anomalocarids. In 2014, it was found that tardigrades were more closely related to arthropods than velvet worms.
  
  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. 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.
  
  In the 1990s, molecular phylogenetic analyses of DNA sequences produced a coherent scheme showing arthropods as members of a superphylum labelled Ecdysozoa ("animals that moult"), which contained nematodes, priapulids and tardigrades but excluded annelids. This was backed up by studies of the anatomy and development of these animals, which showed that many of the features that supported the Articulata hypothesis showed significant differences between annelids and the earliest Panarthropods in their details, and some were hardly present at all in arthropods. This hypothesis groups annelids with molluscs and brachiopods in another superphylum, Lophotrochozoa.
  
  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.