We investigated possible relationships among pseudocoelomates (aschelm
inths) by using cladistic analysis to uncover all most-parsimonious tr
ees from a data set comprising 45 morphological characters of 11 phyla
(3 acoelomate, 7 pseudocoelomate, and 1 coelomate). Analysis of our m
atrix yielded 2 most-parsimonious trees, differing only in relative pl
acement of 3 taxa (Kinorhyncha, Loricifera, and Priapulida). Tree topo
logy suggests that the pseudocoel either evolved twice, or that it evo
lved once and was then independently lost in Gastrotricha and in Gnath
ostomulida. Tree topology also is consistent with derivation of the ps
eudocoel from a coelom. From our analysis and that of other morphologi
cally-based studies, we conclude that there are 2 main lines of pseudo
coelomate evolution: (1) a minor clade (Acanthocephala + Rotifera) def
ined principally by a syncytial epidermis, intracytoplasmic lamina, an
d sperms with anterior flagella; (2) a major clade (the 5 remaining ph
yla), defined primarily by a cuticle that is molted. Within the major
clade are 2 subclades (Nematoda + Nematomorpha; Kinorhyncha + Loricife
ra + Priapulida). Using morphological data, the chief issues remaining
to be resolved are placement of Gastrotricha and Gnathostomulida, and
evolution of the pseudocoel. No analysis based solely on morphologica
l data has so far produced a tree congruent with those based on molecu
lar data. Nevertheless, there is a general consensus that (1) the pseu
docoel is of little phylogenetic significance and (2) the pseudocoelom
ates are probably polyphyletic.