Sauropod nostrils: Where were they?

Short answer:
For whatever reason, derived sauropods shifted the external naris away from the mouth. It would appear illogical to extend soft nostrils back close to the mouth, as Witmer 2001 proposes, over the exterior of the maxillary basin (Fig. 1), which varies greatly (Fig. 2).

Witmer 2001 proposed an anterior nostril position
within the nasal basin anterior to the bony external naris in sauropods (positions A and B in Fig. 1, green dot in Fig. 2) and a similar anterior position in other dinosaurs based on an anterior position in most lepidosaurs, crocs and birds. In every photo example presented by Witmer the nostril forms only a small opening relative to the bony external naris.

Witmer 2001 also provided several exceptions to that pattern:

1. “Cormorant (Phalacrocorax) simply lacked a ßeshy nostril altogether (a diving adaptation)
2. The bony nostril of geckos is so small that the fleshy nostril occupied almost its entire extent.
3. The most significant exception was among monitor lizards (Varanus). Some species (e.g., V. griseus, V. dumerili, V. exanthematicus) have a fleshy nostril located in the middle to caudal half of the much enlarged bony nostril.”
4. Witmer concludes: “Given the diversity of amniotes, one would expect to find additional exceptions.”

As everyone knows,
all tetrapods are capable of inhaling and exhaling through the mouth, which becomes important in panting for internal cooling and when exercise requires more oxygen. The external naris is principally for olfaction and the anterior position of the nostril within the naris maximizes the amount of soft tissue that can be exposed to incoming odors and pheromones.

A tracing of the external and internal nares in sauropods
(Fig. 2) and a simplified guess connecting the two in lateral view, shows

1. the elevation of the external naris (pink) relative to the internal naris (blue)
2. the spacious airway (blue) in sauropod skulls.
3. the reduced airway proposed by Witmer (green) if skin extended the external naris to the anterior nasal basin
4. the easy drainage of rainwater if allowed to directly enter the nostrils (pink) in sauropods (probably unimportant, but thought I’d mention it since most nostrils/nares, except whales and crocs, are anterior to lateral, not dorsal)
5. When dipping the head down to drink, the internal naris were able to fill with water that drained into the throat whenever the lips were sealed and the skull was elevated. That is marginally different from the ostrich drinking behavior (below).
6. Based on the ostrich example, the sauropod nostril may have extended from 1/3 to 2/3 the area of the external naris in brachiosaurs, to the entire naris in the relatively small external naris of Diplodocus (Fig. 2).

Witmer 2012 (YouTube video below)
provided an ostrich skull in which tissue labeled ‘airway’ completely filled the external naris.

Unfortunately,
the Witmer video does not show the nostril seen in an ostrich photo (Fig. 3). Confusing. That should have been somehow clarified, because the nostril is present in vivo, not in the µCT scan.

The small head on the end of a long neck
of an ostrich is analogous to the small head and long neck of sauropods when it comes to breathing and drinking. In the ostrich the nostril is one third the size of the naris and located within the naris, more or less anteriorly. Drinking would have been similarly done, with similar problems to get over, like transferring a throat-full or snout-full of water to the stomach by elevating the head and neck.

In a future post

we’ll look, from a scientist’s perspective, why scientists shy away from attempting to replicate discoveries. On the other hand, I revel in testing published hypotheses because so often they leave their work unfinished or misguided one way or another. All the loose ends need to be tidied up.

References
Witmer LM 2001. Nostril position in dinosaurs and other vertebrates and its significance for nasal function. Science 293, 850-853. PDF