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Crustacea and arthropod relationships. Festschrift for Frederick R. Schram
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In , Schram, F. In the following, we discuss examples how a pan-monophylum concept could be applied to crustaceans. If we would now apply the pan-monophylum concept s as proposed, it would not include Hexapoda, as Hexapoda has extant representatives. Hexapoda would in this case represent the sister group to Pan-Crustacea or Pancrustacea.
Figure 1: Different constructions of pan-monophyla based on different phylogenies. The term Pancrustacea is therefore ambiguous if used in this way. If one wants to stress this specific supposed sister-group relationship between Eu- Crustacea and Hexapoda, the non-ambiguous term Tetraconata is available [ 19 ].
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Example 2 Hexapoda as a crustacean in group: Different authors have used Pancrustacea as a substitute for Eu- Crustacea, but with an emphasis on Hexapoda being a deeper ingroup of it. In this way, the term Pancrustacea is in fact equivalent with Eucrustacea Figure 1C. Eucrustacea has been characterised based on autapomorphies [ 17 ].
Hence, adding a group somewhere inside does not change the validity of this group. As for Reptilia it has now been stressed for many years that Crustacea is not valid, but a paraphylum.
Crustacean and Arthropod Relationships
Yet, Reptilia has been re-established as simply including birds and is now a valid monophyletic group [ 20 ]. Similarly, if Hexapoda is indeed an ingroup of Eucrustacea, this would simply mean that hexapods are eucrustaceans. Concluding: Pancrustacea in this use differs to that of other applications, has no relation to any pan-monophylum concept, as it does not make a reference to fossils and is simply equivalent to Eucrustacea. We generally face the problem that the exact relationships of Eucrustacea are partly unclear.
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The exclusively fossil Phosphatocopina is closely related to Eucrustacea, based on numerous shared characters of the feeding apparatus differentiated hypostome-labrum complex, sternum with paragnaths, different details such as setation [ 21 ]. Yet, most of these, if not all characters appear to be present also in Myriapoda and Hexapoda, or at least these groups can be deviated from an ancestor with such morphology [ 22 - 24 ].
Hence, we cannot reliably resolve Hexapoda, Myriapoda, Phosphatocopina and Eucrustacea. Still, we can think of the different possibilities. For example, if Hexapoda would be definitely identified as a stable ingroup of Eucrustacea, 'Pancrustacea' could be applied, referring to Eucrustacea plus Phosphatocopina but excluding the next branch with extant representatives which could be Myriapoda.
In this case, 'Pancrustacea' would comprise Eucrustacea with Hexapoda as an ingroup plus Phosphatocopina.
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Hence 'Pancrustacea' would be synonymous to Labrophora Figure 1B ; as characterised by Siveter et al. Yet, it is possible that Myriapoda is more closely related to Eucrustacea with Hexapoda as an ingroup than Phosphatocopina to Eucrustacea. In this case, the pan-monophylum concept could not be applied as there would be no known fossil branching off the evolutionary lineage towards Eucrustacea after the branch of Myriapoda. The same scenarios can also be applied if Hexapoda and Myriapoda form a monophyletic Tracheata.
And an alternative name is instead available. Crustacea sensu lato includes also a number of fossil representatives which have been addressed as stem-crustaceans [ 18 , 28 ] or stem-mandibulates [ 28 ], better as early derivatives of the evolutionary lineage towards Eucrustacea [ 18 , 21 , 29 , 30 ]. There is no application of the pan-monophylum concept that can be used as an equivalent for Crustacea sensu lato.
This is due to the fact that the sister group of Crustacea sensu lato, Agnostina, has no extant representatives and therefore would have to be included into any pan-monophylum also including the early representatives of Crustacea sensu lato together with many further groups, see below. If we now apply the pan-monophylum concept based on Crustacea sensu lato or on a monophyletic group Eucrustacea that includes Hexapoda and Myriapoda [ 31 ], 'Pancrustacea' would be a very large group including also Agnostina, probably trilobites, possibly also nectaspidids and many other fossil euarthropods Figure 1D [ 32 ] as the next group with extant representatives is Chelicerata sensu lato [ 33 , 34 ].
Thus, 'Pancrustacea' in this case would equalize that part of Euarthropoda that does not comprise chelicerates[ 35 ]. For the different possible meanings other less ambiguous names are available. With the term 'Panarthropoda' we face similar problems as with 'Pancrustacea'. Currently, as for 'Crustacea' there is no generally well-recognised monophyletic group named 'Arthropoda' exceptions see below. Thus, we have to discuss several cases independently. As Onychophora and Tardigrada are groups with numerous extant representatives, this use has to differ from any construction of a pan-monophylum.
Hence, the term is ambiguous in this case. The most stable use of Euarthropoda is probably the one introduced by Walossek [ 17 ] as it is again an autapomorphy-based characterisation, founded on 1 a head including five segments, the ocular and four appendage-bearing ones, and 2 the formation of a basipod on the appendages of the second post-ocular segment and further posterior ones although also here it would be more stable to choose one of the two, i. Based on this use of Euarthropoda, 'Panarthropoda' could, for example, include all fossil representatives down before Pentastomida, Tardigrada or Onychophora [ 25 ] branch off the evolutionary lineage towards Euarthropoda.
This use of 'Panarthropoda' could equalise more or less the monophyletic group Arthropoda sensu stricto, also an autapomorphy-based group including all sclerotised arthropods Figure 1F [ 36 ]. Yet, this application of 'Panarthropoda' to Euarthropoda most likely additionally includes the one or other lobopodian [ 37 ].