About this edition. This is Antoni Milewski’s post, republished with his permission. Only the photographs he chose are used. Those on iNaturalist under a Creative Commons licence are shown; the rest are hosted externally and are given as links. No images have been substituted. Read the original on iNaturalist.

The eyes and ear pinnae of ungulates vary in placement, according to habitat and anti-predator strategy.

In the case of the eyes, there is variation in how far to the sides, and how high on the head, they are placed.

In the case of the ear pinnae, there is variation in how high on the head they are placed, and in how they are oriented (upright, sideways, or downwards).

Eyes

An extreme example of lateral placement of the eyes is the pronghorn (Antilocapra americana):

CitedAlamy·henrysforkwildlifealliance.org·sdakotabirds.com

The opposite extreme is Myotragus balearicus (Wikipedia), which evolved on small islands, virtually free of predators:

CitedFlickr·Twitter 1·Wikimedia 1·Twitter 2·Wikimedia 2

Turning to the placement of the eyes high on the head:

A peculiar example is Hippopotamus amphibius:

CitedAlamy

The extinct Hippopotamus gorgops was even more specialised:

CitedWikipedia·quoracdn.net

However, there is a similar configuration in warthogs (Phacochoerus spp.), which are the suiforms with the least affinity for water:

nataliefieldPaulshof · 2018-01-16CC BY-NCObservation

Warthogs live on land, but have a short neck and legs

CitedDreamstime

and kneel while foraging:

almut_66Ngorongoro Crater · 2017-09-24CC BY-NCObservation

Thus, they risk not spotting danger above the grass.

CitedDreamstime 1·Shutterstock·Dreamstime 2·Dreamstime 3·Dreamstime 4·Dreamstime 5·Dreamstime 6·Dreamstime 7

tonyrebeloH6 Satara-Nwanetsi Rd · 2021-06-07CC BY-SAObservation

In their own way, warthogs are unique in having the eyes placed above the ‘horns’:

CitedDreamstime

Ear pinnae

Among wild spp., an extreme example of low placement is Syncerus, in which the ear pinnae are shaded by the horns:

Citedkidadl.com·mediastorehouse.com.au·Wikimedia

However, the most extreme species is a domestic one, viz. Bos indicus:

Cited123rf.com

In large bovines, the function of the ear pinnae seems to be for thermoregulation as much as hearing (see the following):

CitedRelated post

Applying this framework to grey rhebok

The grey rhebok (Pelea capreolus) emerges as having

  • possibly the most upright ear pinnae of any ungulate,
  • unusually – albeit not extremely – high-placed eyes, and
  • eyes as laterally-placed as in any bovid.

Please see the following:

hhodgsonVergelegen Wine Estate · 2022-10-30CC BY-NCObservation
leejonesCape Point · 2016-04-23CC BY-NCObservation

How can the combination of high eyes and extremely upright ear pinnae in the grey rhebok be explained, in terms of adaptation?

It is true that the habitat of the grey rhebok is generally open, with low vegetation. This would seem to make high placement/orientation of the sense-organs redundant.

richardadcockWaboomberg · 2016-11-06CC BY-NCObservation

Furthermore, the grey rhebok partly coexists with Damaliscus pygargus pygargus, one of the most conspicuous of bovids, presumably in adaptation to open environments.

tonyrebeloDe Hoop Nature Reserve · 2015-12-25CC BY-SAObservation

However, an apparent paradox is that the grey rhebok – in complete contrast to D. p. pygargus – has extremely inconspicuous (cryptic) colouration:

CitedRelated post

garthbushellGalway Amajuba · 2021-10-04CC BY-NCObservation

This is partly explained by the facts that the grey rhebok is one of the few ungulates that penetrates mature stands of fynbos:

lallenWest Coast DC · 2016-08-24CC0Observation

has a body size such that the head of the standing figure corresponds approximately to the height of mature fynbos:

gawieBreede River DC · 2018-04-16CC BY-NCObservation
thibaudaronsonBontebok NP · 2016-03-28CC BY-SAObservation
tonyrebeloMaajiesriver Cedarberg · 2013-11-23CC BY-SAObservation

and differs from reedbucks (Redunca arundinum

lemonculRumphi · 2020-08-20CC BY-NCObservation
magdastluciaiSimangaliso · 2021-10-24CC BY-NCObservation
mortenCKGR · 2019-10-22CC BYObservation

and Redunca redunca) in not lowering the head, and partly turning the ear pinnae back, when initially alarmed:

Citedstock.adobe.com

The combination of features peculiar to the grey rhebok can be summarised as follows, relative to reedbucks.

The eyes are as high on the head, and as laterally placed, as in Redunca fulvorufula.

Compare grey rhebok with R. fulvorufula:

🔒

All rights reserved

skhokho · WC · 2022-04-23

Also cited on iNaturalist →

CitedAlamy

However, the eyes are not as high-placed as in Redunca arundinum

joachimKasungu · 2013-08-05CC BY-NC-NDObservation
magdastluciaWestern Shores · 2017-07-22CC BY-NCObservation
happyasacupcakeIthala GR · 2018-02-14CC BY-NCObservation
bartwurstenPicada 4 near junction 3 · 2015-12-08CC BY-NCObservation
happyasacupcakeIthala GR · 2019-01-14CC BY-NCObservation
jouberthEhlanzeni · 2018-07-26CC BY-NC-NDObservation

and possibly Redunca redunca:

joachimKatavi · 2015-05-30CC BY-NC-NDObservation
pgtzAmboseli National Park · 2022-07-05CC BY-NCObservation

The ear pinnae are far more upright in the grey rhebok than in any species of reedbuck.

Compare grey rhebok

gigilaidlerCape of Good Hope Nature Reserve · 2022-04-30CC BY-NCObservation
nicolaasmyburghSwellendam · 2022-01-09CC BY-NCObservation

with R. arundinum

magdastluciaiSimangaliso · 2021-07-04CC BY-NCObservation
happyasacupcakeTshokwane-Lower Sabie rd · 2019-12-04CC BY-NCObservation

and R. redunca:

martingrimmBagamoyo · 2018-06-02CC BY-NCObservation
🔒

All rights reserved

lecomte · A7 · 2018-01-24

Also cited on iNaturalist →

I suggest that this peculiar combination is consistent with the habitats and behaviour peculiar to the grey rhebok. Whereas reedbucks tend to be adapted to the tall grass of seasonal marshes, the grey rhebok is dually adapted to mature fynbos and short vegetation.

CitedRelated post

Addenda from the thread of comments

The author added the following in comments beneath the original post.

The following shows the position to which the ear pinnae of the grey rhebok sink in death, before rigor mortis:

Citedafricahunting.com

The upright orientation of the ear pinnae in the grey rhebok has sometimes not been realised by taxidermists:

Citedthetaxidermystore.com·eBay

An auricular semet is a subtle pattern of colouration on or adjacent to the ear pinnae, functioning for intraspecific (social) communication. It is activated by movement, particularly the swivelling of the ear pinnae according to mood/emotion and the direction of attention/intention.

The largest-scale patterns of colouration on/near the ear pinnae may qualify instead as auricular flags, because – once activated by movement – they are capable of making the whole figure conspicuous, including to scanning predators.

An unambivalent example of an auricular semet occurs in Axis axis:

CitedDreamstime 1·Fine Art America·rlbennettphotography.com·Freepik 1·Freepik 2·Bigstock·agefotostock·Dreamstime 2·Dreamstime 3

If you focus on the dorsal surface of the anterior base of the ear pinna, you may notice a crisp dark/pale (actually black/white) pattern, on a scale within centimetres.

The configuration changes depending on the position of the ear pinna relative to the head. This means that, as the ear pinnae move in vigilance, attentiveness, or emotion, the movement is accentuated by the pattern of dark/pale contrast.

This pattern cannot be dismissed as functioning mainly for inconspicuousness in conformation with the general spotted pattern of the animal. This is because a) camouflage – which is by definition stationary – cannot explain why the pattern has been configured to change so much with the movements of the ear pinnae, and b) for inconspicuousness, the simplest pattern would be plainness of the whole ear pinna and its base, without any pattern at all:

CitedDreamstime

Other spp. of cervids have different auricular semets, e.g. Cervus nippon:

CitedGetty Images

Please see also adjacent Posts:

CitedRelated post

Compare Blastocerus dichotomus with Axis axis:

CitedShutterstock

A further observation, cited without comment:

Credits. Photographs reproduced from iNaturalist are shown under the Creative Commons licence stated in each caption and remain the copyright of the observers named. All other images cited by the author are hosted externally and are linked, not reproduced.

By Antoni V Milewski and Anthony J Mills. Text by Antoni V Milewski, posted on iNaturalist 2 December 2022 and republished here with permission. Read the original.