The Systematic Relationships of the Monotremata: The Ontogenetic Evidence
The Systematic Relationships of the Monotremata: The Ontogenetic Evidence
Ulrich Zeller
Museum f,r Naturkunde, Institut f,r Systematische Zoologie,
Invalidenstrasse 43, 10115 Berlin, Germany
Modern systematics must be based on a holistic approach
to the organism. This includes not only molecular genetics
and its links to morphogenesis, but also morphogenesis
by itself has to be considered. In this context the
systematic relations of monotremes are reevaluated.
The Monotremata and the other Recent Mammalia (Theria)
as well as all other groups of Mesozoic mammals share
a number of synapomorphies in comparison with their
synapsid ancestors that characterize them as a monophyletic
group (e.g. squamoso-dentary jaw joint as secondary
jaw joint; caudal part of the lower jaw functionally
separated from the jaw apparatus transmitting air-born
sound; ethmoideal region with extended peripheral olfactory
organ, recessus ethmoturbinalis and lamina cribrosa;
integument rich in glands including lacteal gland;
reproduction characterized by the combination of parental
care, lactation, yolk reduction and fetomaternal metabolism
(placentation). Furthermore, each of the three Recent
mammalian groups (Monotremata, Marsupialia, Placentalia)
shows its own mosaic of retained primitive features
(plesiomorphies) and derived features (apomorphies).
The Monotremata have retained a series of plesiomorphic
characters of the last common ancestor of Recent mammals
which were lost in the Theria (e.g. oviparity, cloaca,
shoulder girdle, egg tooth, septomaxillary, ectopterygoid,
temporal canal, pila antotica, cartilago scleralis).
On the other hand, Theria have incorporated plesiomorphic
features from their common ancestor which were lost
in Monotremata (e.g. pila metoptica, lacrmimal, parasphenoid).
Monotremata and Marsuplialia share plesiomorphic mammalian
characters, such as meroblastic cleavage, tertiary
egg membrancs, precocial offspring, short development
within fetal membranes and extensive lactation - Monotremata
are autapomorphic with regard to the formation of the
secondary side wall of the braincase (lamina obturans).
In addition, Marsupialia display autapomorphic characters,
e.g. reduction of alantois and mainly choriovitelline
placentation, the loss of pila postoptica and the inflection
of the angular process of the dentary. The Eutheria
are autapomorphic in having achieved a feto-maternal
immuno-tolerance as a prerequisite of a long intrauterine
development and intensive placentation. Marsupialia
and Placentalia are synapomorphic, for example, in
the formation of the secondary side wall of the braincase
by the alisphenoid. The individual autapomorphies of
Monotremata and Theria suggest an early dichotomy of
the phyletic lines leading to the Monotremata and the
Theria respectively. There is no developmental synapomorphy
of Monotremata and Marsupialia suggesting the taxon
Marsupionta. Instead, the sister group relationship
of monotremes and therians is well established.