Jn. Caira et al., On the phylogenetic relationships among tetraphyllidean, lecanicephalideanand diphyllidean tapeworm genera, SYST PARAS, 42(2), 1999, pp. 77-151
This study had two main objectives: (1) to construct an extensive, explicit
list of characters and character states that might serve as a starting poi
nt, and perhaps even a model, for the compilation of a more complete list o
f characters for all cestode taxa; and (2) to use this character list to ge
nerate a hypothesis of the phylogenetic relationships among species represe
nting most of the tetraphyllidean, lecanicephalidean and diphyllidean gener
a. Specimens of one species in each of 48 genera of tetraphyllideans, eight
genera of lecanicephalideans, the three genera of diphyllideans, two gener
a of proteocephalideans and two genera of trypanorhynchs, were examined as
whole-mounts and sections, with light and scanning electron microscopy. A l
ist of 120 morphological characters was compiled. Four phylogenetic analyse
s were conducted using PAUP* and/or NONA. The first was a comprehensive ana
lysis with the 56 tetraphyllidean and lecanicephalidean species as ingroups
and the remaining seven species as outgroups. The second was an analysis o
f the three diphyllidean species as ingroups and the two proteocephalidean
and the two trypanorhynch species as outgroups. The third was an analysis o
f the eight lecanicephalidean species and the "tetraphyllideans" Echeneibot
hrium sp. and Pseudanthobothrium n. sp. as ingroups and an outgroup consist
ing of the seven species used as outgroups in the first analysis. In the fo
urth analysis, the ingroup consisted of the 14 hooked tetraphyllideans (onc
hobothriids), and the outgroup consisted of the seven species used as outgr
oups in the first analysis. The results of these analyses support the follo
wing phylogenetic hypotheses: The diphyllideans are monophyletic and Echino
bothrium n. sp, and Macrobothridium sp. are more closely related to one ano
ther than either is to Ditrachybothridium macrocephalum. The tetraphyllidea
ns, lecanicephalideans and proteocephalideans are more closely related to e
ach other than they are to the diphyllideans or the trypanorhynchs. The ord
inal status of the lecanicephalideans is dubious. The lecanicephalidean spe
cies are more closely related to some of the tetraphyllidean taxa than thes
e tetraphyllidean taxa are to the remainder of the tetraphyllidean tare. Th
e proteocephalideans appear to belong within the tetraphyllidean clade. The
"tetraphyllidean" species Echeneibothrium sp. and Pseudanthobothrium n. sp
. are members of the lecanicephalidean clade. The position of "Discobothriu
m" n. sp. within the lecanicephalideans is dubious. Within the tetraphyllid
eans, the non-acetabulate species Litobothrium daileyi, Disculiceps galapag
oensis and Cathetocephalus sp. are the most basal members of the group. The
family Onchobothriidae is monophyletic, as it is currently defined. Within
the onchobothriids, the uniloculate species are basal to the multiloculate
species; the species with unipronged hooks are basal to the species with m
ultipronged hooks. Although relationships among the phyllobothriids. as the
y are currently defined, remain poorly resolved, the family Phyllobothriida
e is not monophyletic, These results suggest that some aspects of the class
ification of the lecanicephalidean and tetraphyllidean taxa require revisio
n. However, such revision should be based on further analyses including a b
roader representation of the genera and species in these groups.