D. Walossek et Kj. Muller, PENTASTOMID PARASITES FROM THE LOWER PALEOZOIC OF SWEDEN, Transactions of the Royal Society of Edinburgh. Earth sciences, 85, 1994, pp. 1-37
Small phosphatised fossils from late Cambrian limestones of Vastergotl
and, Sweden, share major external features with larval extant Pentasto
mida, such as a prominent head with two pairs of stumpy limbs adapted
for attachment, and a slender trunk of four portions. Even such detail
s, as paired forehead structures, pores on the inner edges of the head
limbs and paired papillae at the rear of the trunk correspond with st
ructures of extant pentastomid larvae. Neither the fossils nor the Rec
ent pentastomids add any additional body segments during growth (segme
nt constancy). Since characters of this kind and in this combination d
o not occur elsewhere, the fossils are recognised as true Pentastomida
. Major differences, such as distinctly divided head limbs, partial oc
currence of vestigial trunk limbs, and a different mode of trunk devel
opment during growth can be explained as representing merely the plesi
omorphic state of characters of Pentastomida, indicating that the foss
ils are representatives of its stem-group prior to branching into the
two Recent lineages. The fossils clearly document the marine origin of
the Pentastomida, and that their specific morphology and parasitic li
fe style were already established in the late Cambrian at a high degre
e of diversification, long before the terrestrialisation of their pres
ent final hosts, the tetrapods. General arthropod affinities are recog
nisable not least in the nature of the limbs, but the morphology of st
em- and crown-group pentastomids gives no clues for closer relationshi
p with any of the major (eu)arthropod taxa.