R. Frey et Rr. Hofmann, EVOLUTIONARY MORPHOLOGY OF THE PROBOSCIDEAL NOSE OF GUENTHER DIKDIK (RHYNCHOTRAGUS-GUENTHERI THOMAS, 1894) (MAMMALIA, BOVIDAE), Zoologischer Anzeiger, 235(1-2), 1996, pp. 31-51
The internal structure of the conspicuous proboscis of three male Guen
ther's dikdiks (Rhynchotragus guentheri) was investigated. The probosc
is consisting of connective tissue and musculature is attached to the
rostral part of the head. This head region is characterized by a reduc
tion and caudal displacement of bony and cartilaginous components. The
ventral concha in particular is small and reduced in length. The leng
th of the nose amounts to 60-70% of skull length. Taking into account
the anatomically-determined curvature the length of the air passage th
rough the nose may reach 80% of skull length. Proboscis tubes are supp
orted by a muscular pad originating from the incisive bone. Evolution
of the proboscis tubes caused a rostral separation of the ventral nasa
l meatus from both the middle and dorsal nasal meatus. The three passa
ges remain separated up to the caudal end of the proboscis tubes. The
structure of the respiratory region is conspicuously different from a
typical bovid nose. It consists of an umbrella-shaped cartilage with i
ts concave surface rostrally oriented and a posterior cartilaginous ch
amber the interior of which is accessible only through a small medial
opening. A thick bundle of longitudinally-oriented, anastomosing, thic
k-walled veins passes along the dorsal surface. In addition, larger lo
ngitudinally-oriented fascicles of veins run along the ventral surface
of the cartilaginous chamber, along the medial side of the dorsal con
cha, and within the plica recta; all of these vessels underlie the del
icate nasal mucosa. The longitudinal fascicles of veins facilitate a c
ountercurrent exchange between air stream and blood stream only during
the expiratory phase. During inspiration, air and blood flow in the s
ame direction. The rostral opening of the nasolacrimal duct is situate
d at the medial side of the cartilaginous umbrella. Below its mucosa t
he nasal septum is equipped with a rostral and a caudal cavernous body
each consisting of numerous longitudinally-oriented veins. The caudal
one extends laterally into the opening of the cartilaginous chamber.
The relevance of these anatomical results with respect to thermoregula
tion and water conservation is discussed using physiological data from
the literature. At ambient temperatures below 30 degrees C (either at
night or in the shade) the exchange functions of the nose are employe
d to recover effectively water. At ambient temperatures above 30 degre
es C (during daytime in the sun) the evaporative cooling functions of
the nose (and of the oral cavity) have to be employed to a greater ext
ent accompanied by an increased loss of water. Water shortage causes a
partial dehydration of the large amount of mucus covering the nasal m
ucous membrane surfaces. This might allow the recovery of water vapour
during exhalation. Fat deposits in honeycomb-shaped cavities of the s
kull base probably serve to insulate the complex system of blood vesse
ls consisting of the carotid rete (= rete mirabile epidurale) and its
surrounding venous network (= sinus cavernosus). The cartilaginous ele
ments of the respiratory region and the accompanying mucous membrane f
olds as well as the contiguous muscles can be clearly homologized with
structural elements of a typical bovid nose. In the evolution of the
proboscideal nose, rostrally located parts became reduced in length an
d were displaced caudally. This resulted in the cartilaginous elements
of the lateral nasal wall and of the nostril, together with the fasci
cles of veins, undergoing change to a thermoregulative function (heat
exchange, water recovery). The proboscis tubes lined with cutaneous mu
cosa and the circular nostrils directly in contact with each other and
pointing downward are recently evolved structures. Presumably they im
prove the insulation of the vapour exchanger. Of all the Madoquini, R.
guentheri lives in the most arid habitats and is largely independent
of drinking water. Thus, the development of a proboscideal nose and th
e transformation of the respiratory region have evolved as adaptations
for the enhancement of water recovery.