B. Benslimane et al., TRICHOSTRONGYLINA, NEMATODE PARASITES OF AMPHIBIANS AND REPTILES - TAXONOMIC, PHYLOGENETIC AND BIOGEOGRAPHIC PROBLEMS, Systematic parasitology, 35(3), 1996, pp. 179-206
Except for four species of the Mertensinematinae (Molineoidea) not con
sidered in this work, the Trichostrongylina parasitic in amphibians an
d reptiles, comprising 15 genera and 105 species, are reviewed. The mo
rphological characteristics of each species are analysed. The most imp
ortant characteristics are provided by the disposition of the caudal b
ursal rays, the morphology of the synlophe (at the oesophago-intestina
l junction and mid-body level) and by the anatomy of the spicules. The
species are classified into seven groups: Group 1: ''relict'' species
with six lips and well-developed buccal capsule (six genera and 16 sp
ecies). Group 2: ''ancient'' species with one or two primitive charact
ers (eight genera and 15 species). Seventy-four species, without primi
tive characters, comprise the genus Oswaldocruzia which is subdivided
into five groups. Group 3: Oriento-Ethiopian species with non-''idiomo
rphic'' spicules with two to three tips (10 species). Group 4: Neo-Eth
iopian species with non-''idiomorphic'' spicules with numerous tips (1
1 species). Group 5: Holarctic species with ''idiomorphic'' spicules w
ith the spicular fork divided above the distal third of the spicule le
ngth (24 species). Group 6: Continental Neotropical species with ''idi
omorphic'' spicules and spicular fork divided within the distal third
of the spicule length (21 species).Group 7: Caribbean Neotropical spec
ies with modified ''idiomorphic'' spicules, the three main branches of
which are each divided into numerous tips (eight species). The geogra
phical distribution of the species appears to be of greater significan
ce than the host spectrum and suggests the following biogeographical h
ypotheses: Group I, entirely Gondwanan, diversified during the Cretace
ous era prior to the separation of the southern continents. The expans
ion of Group 2, which is represented by Gondwanan and Oriental species
mainly from India, Malaysia and Indochina, could have occurred throug
hout South-East Asia at the end of the Cretaceous era when India colli
ded with Eurasia. Group 3 could be interpreted either as a migration f
rom Asia to Africa during the upper Eocene, or more likely during the
Miocene, or, by a dispersion due to the migrations of Bufo melanostict
us. Group 4 may result from the expansion of the former group in the E
thiopian region. Group 5 could be interpreted as a colonisation of wes
tern Europe and the Nearctic from Asia during the Tertiary. Group 6 co
uld have arisen after the migration of the Neartic species to the Neot
ropical region during the Pliocene period and Group 7 by the expansion
of the former group in the Caribbean. Amongst the Trichostrongylina,
the ''relict'' and ''ancient'' genera parasitic in amphibians and rept
iles can only be compared with the genera parasitic in birds and mamma
ls, probably dating from the Palaeocene period (mainly marsupials and
primitive insectivores). On the other hand, the genus Oswaldocruzia ca
n be compared with other members of the Molineoidea parasitic in fissi
peds, Pholidota and Chiroptera dating from a later period (Eocene) (se
e Durette-Desset, 1971). Thus, the Trichostrongylina of amphibians and
reptiles are distinguished by the persistence of very old species, th
e evolution and the geographical distribution of which was arrested fo
r millions of years, and by the existence (in some regions) of groups
undergoing full evolutionary expansion. The new taxa proposed in the p
aper are: Bakeria (Moravec & Sey, 1986) status emend. (= Bakeria Morav
ec & Sey sub. g.), Ragenema n. g., Ragenema robustum (Baker, 1982) n.
comb. (= Oswaldocruzia robusta), Typhlopsia quentini (Durette-Desset,
1980) n. comb. (= Trichoskrjabinia quentini), T. secundus (Pinnell & S
chmidt, 1977) n. comb. (= Trichoskrjabinia secundus), T. gansi (Crusz
& Ching, 1975) n. comb. (= Oswaldocruzia gansi), T. limnodynastes (Joh
nston Bi Simpson, 1942) n, comb. (= Oswaldocruzia limnodynastes), T. l
egendrei (Chabaud & Brygoo, 1962) n. comb. (= Oswaldocruzia legendrei)
.