THE JOURNAL OF EXPERIMENTAL ZOOLOGY 282:507–525 (1998)
© 1998 WILEY-LISS, INC.
Survey of the Oviduct of Salamandrids With Special
Reference to the Viviparous Species
HARTMUT GREVEN*
Institut für Zoomorphologie und Zellbiologie der Heinrich-Heine-Universität
Düsseldorf, D-40225 Düsseldorf, Germany
ABSTRACT Urodeles include oviparous and a few obligately viviparous species that belong to
one family, the Salamandridae. Oviducts of both groups have basically the same design, but some
modifications became necessary as viviparous species evolved. The entire oviduct of urodeles is
lined by a monolayered epithelium, which is regionally differentiated into large glands and smaller
secretory cells rich in glycoproteins. Secretory products important for fertilization form the egg
jelly, which also is present in viviparous species. In the latter species, however, there appear to be
fewer oviductal convolutions and subdivisions of the glandular part of the organ, as well as fewer
gland cells. Viscous, acidic secretory products predominate in viviparous species, whereas resis-
tant neutral mucins predominate in oviparous species. In viviparous species, the caudal portion of
the oviduct became altered to retain the developing offspring. This “uterus” lacks glands, but
epithelial cells produce carbohydrate-rich material. Offspring remain in the uterus anywhere from
several months to five years, depending on the species and climatic conditions. The Na
+
-K
+
-AT-
Pase system, common in vertebrate epithelia, is used to regulate the intrauterine milieu. Subepi-
thelial vessels mediate gas exchange and the removal of waste products, mainly urea. Secretions
of the uterine epithelium may contribute to the fluid surrounding the young, but do not seem to
support nutrition. Sources of nutrition for the young vary from one (sub)species to the next. Some
feed on yolk reserves (Salamandra salamandra and certain of its subspecies), others on unfertil-
ized eggs and siblings (other S. salamandra subspecies and Mertensiella luschani with subspe-
cies), and yet others on degrading embryotrophic eggs as well as on cells derived from a specialized
uterine trophic zone (S. atra and subspecies). Certain basic alterations in the uterine epithelium
during pregnancy are most likely under endocrine control. Others, such as the flattening of cells,
the discharge of secretory products, and the development of the trophic zone, may be induced by
interactions with the offspring. J. Exp. Zool. 282:507525, 1998. © 1998 Wiley-Liss, Inc.
Viviparity, a mode of reproduction in which young
are carried in a parent’s body during a major por-
tion or during the entire period of development, has
evolved many times within the vertebrate phylum
(for discussions of definitions: Wourms et al., ’88;
Wake and Roth, ’89; Wake, ’93). Viviparous species
are found in all three extant orders of Amphibia,
most of them among the Gymnophiona, and only a
few among Anura and Urodela (Wake, ’93). Of the
urodeles, only members of the family Salaman-
dridae, i.e. all species of the Palaearctic genus
Salamandra and the related species Mertensiella
luschani are obligately viviparous. The exact
number of species and/or subspecies belonging to
these genera is still a matter of debate (references
in Greven and Thiesmeier, ’94). Larvae at diverse
stages of development, or even metamorphosed
young, are born in Salamandra species, but
Mertensiella luschani gives birth only to trans-
formed offspring. Embryos and larva develop in a
modified caudal portion of the Müllerian ducts (=
oviducts). Usually it is called a uterus, a term that
is justified because the oviducts of most verte-
brates are homologous and have similar functions.
Nutritive support of the young during development
ranges from the initial yolk stores to oophagy,
adelphophagy, and even epitheliophagy (for refer-
ences see Greven and Thiesmeier, ’94).
The general bauplan and basic functions of the
oviducts in viviparous salamandrids are largely
similar to those of oviparous species. Neverthe-
less, viviparity and the trophic patterns mentioned
above, i.e. lecithotrophy (nutrients provided only
*Correspondence to: Hartmut Greven, Institut für Zoomorphologie
und Zellbiologie der Heinrich-Heine-Universität Düsseldorf, Univer-
sitätsstrasse 1, D-40225 Düsseldorf, Germany.