R. Frey, COPULATORY ORGAN, MATING POSTURE AND LOCO MOTION - THEIR INTERRELATIONSHIP IN NONMAMMALIAN VERTEBRATES, Journal of zoological systematics and evolutionary research, 33(1), 1995, pp. 17-31
Copulatory organ, mating posture and locomotion: their interrelationsh
ip in non-mammalian vertebrates. This paper demonstrates, for the vert
ebrate groups discussed, a relationship between mating posture, the le
ngth of the copulatory organ, and the respective modes of locomotion.
The modes of locomotion in the majority of non-mammalian vertebrate gr
oups presuppose a massive, heavily muscularized, and relatively long t
ail, which may either reinforce lateral flexions of the trunk or (in a
nimals with rigid trunk) act as the only means of propulsion (e.g. in
Chondrichthyes, Teleostei, Lepidosauria, and Crocodylia). Such a massi
ve tail with gradual transition to the trunk necessitates a lateral ma
ting position (regardless of whether the copulatory organ is paired or
unpaired). Animals with a flexible trunk will, therefore, generate st
rongly pronounced laterotruncal flexions ('dynamic lateral bending').
A long copulatory organ is not necessary (e.g. the short hemipenes of
Lepidosauria). Animals with a rigid trunk, on the other hand, will not
be capable of copulating unless by means of a relatively long penis o
r a modified mating posture (e.g. Crocodylia). In the course of phylog
eny, whenever a change in the mode of locomotion allows a reduction in
tail length, this will lead to a corresponding change in mating postu
re, with mating then being possible from a posterior position (e.g. Ch
elonia, Aves, as well as Ascaphus among the Anura). The locomotion of
the latter taxa is characterized by a completely or almost completely
rigid trunk. Thus, contact between male and female cloaca cannot be as
sisted by bending movements in the trunk. To compensate for truncal ri
gidity, there is a trend towards linear extension of the penis, or oth
er male copulatory organs (such as mixopterygia or gonopodia), e.g. in
the pelagic Selachians, in Chelonia, and in the flightless ratites. F
or those birds lacking a penis, the only methods of overcoming truncal
rigidity are stationary locomotion (fluttering without forward moveme
nt), having a mobile, swivelling cloacal region (uropygium), and simul
taneous evagination of the proctodeum of both the male and female. Pen
guins, being unable to perform stationary fluttering, resort to a some
what modified mating posture.