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This Post is dedicated to @gigilaidler, whose painstaking photos in Table Mountain National Park and elsewhere have so clearly illuminated the grey rhebok, among other animals.
The grey rhebok (Pelea capreolus) has remarkably inconspicuous colouration (https://www.inaturalist.org/posts/53474-a-comparison-of-adaptive-colouration-between-lookalikes-grey-rhebok-and-mountain-reedbuck#).
However, like various other animals with cryptic or camouflage colouration, it possesses certain features that become conspicuous when activated by movement.
The obvious example, in the grey rhebok, is the white of the ventral surface of the tail. This feature, which becomes extremely noticeable when the animal flees, can be categorised as a caudal flag (https://www.inaturalist.org/journal/milewski/39938-an-easily-overlooked-but-extreme-adaptation-in-the-grey-rhebok#).
However, there is another feature - dark instead of pale - that has previously been overlooked by zoologists.
I refer to what I call a buccal semet, viz. a small-scale feature of dark/pale contrast at the mouth, which is activated by the cud-chewing movements of the jaws (https://www.inaturalist.org/posts/100574-a-new-feature-of-adaptive-colouration-in-the-grey-rhebok-pelea-capreolus-the-buccal-semet#activity_comment_63701963-aaed-462a-b5af-00a3f88dfe6d).
The buccal semet of the grey rhebok consists of dark short pelage on the anterior part of the side of the mandible, between the lower lip and the 'chin' (https://www.inaturalist.org/observations/73018076 and https://thetaxidermystore.com/african-grey-rhebok-shoulder-taxidermy-mount-14576.html and https://uk.inaturalist.org/observations/185412876).
Please focus on the surface below the side of the mouth in the following:
https://www.shutterstock.com/image-photo/one-grey-rhebok-mountain-zebra-national-1061610842
third photo in https://www.inaturalist.org/observations/117351318
https://www.inaturalist.org/observations/16496842
https://www.inaturalist.org/observations/11867209
https://www.inaturalist.org/observations/148981465
https://www.inaturalist.org/observations/242219998
https://www.inaturalist.org/observations/247271379
https://www.inaturalist.org/observations/247162752
https://www.inaturalist.org/observations/242219998
https://www.shutterstock.com/image-photo/grey-rhebuck-pelea-capreolus-natural-habitat-2166233825
https://www.shutterstock.com/image-photo/grey-rhebuck-pelea-capreolus-natural-habitat-2332821615
scroll in https://antelopesg.org/antelopes/
scroll in https://www.ultimateungulate.com/Artiodactyla/Pelea_capreolusReferences.html
DISCUSSION
The buccal semet in the grey rhebok distinguishes it from reedbucks (Reduncini). It supports the assignment of Pelea to its own exclusive tribe, viz. Peleini, within the Bovidae (http://taxonomicon.taxonomy.nl/TaxonTree.aspx?src=957&id=68333 and https://www.nature.com/articles/hdy2013108).
However, this feature of adaptive colouration seems somewhat convergent with that of the unrelated common duiker (Sylvicapra grimmia):
https://www.inaturalist.org/observations/236945215
https://www.inaturalist.org/observations/243638829
https://www.shutterstock.com/image-photo/grey-rhebok-rhebuck-pelea-capreolus-kruger-571314268
https://www.inaturalist.org/observations/249210122
Furthermore, the darkish vertical mark seen in the grey rhebok is vaguely reminiscent of a pattern in various clades of deer (https://www.inaturalist.org/journal/milewski/55804-the-muzzle-ring-as-a-deep-ancestral-marker-in-deer-part-2-the-gape-spot#).
I postulate the following adaptive function of buccal semets in ungulates:
Ruminants spend much time chewing the cud, which interferes with both forms of non-visual vigilance, viz.
To compensate for these sensory detractions, individuals place themselves (usually lying down in sternal procumbency) within sight of each other, facing in various directions.
When any individual spots (or smells or hears) a potential predator, it immediately stops chewing and turns the ear pinnae towards the source of suspicion. It is crucial, for gregarious enhancement of vigilance, that others notice this attention immediately.
Thus, the sensory system of ruminants is programmed with an acute search-image for a particular combination of cessation of small-scale motion plus accompanying initiation of small-scale motion. This combination consists of
The hypothetical function of buccal semets is to enhance this function, by accentuating the visibility of the chewing motion of the jaws. (Certain ruminants also possess auricular semets, viz. patterns of colouration accentuating the motion of the ears, https://stock.adobe.com/video/sambar-deer-rusa-unicolor-phu-khiao-wildlife-sanctuary-thailand-seen-standing-in-the-rain-facing-the-camera-then-turns-its-head-towards-the-right-side-of-the-frame/438778359?prev_url=detail and https://www.inaturalist.org/posts/55694-ten-best-illustrations-of-auricular-semets-in-deer#.)
It is noteworthy that the buccal semet of the grey rhebok is dark, whereas that of the common impala (Aepyceros melampus) is pale (https://www.inaturalist.org/posts/100574-a-new-feature-of-adaptive-colouration-in-the-grey-rhebok-pelea-capreolus-the-buccal-semet#activity_comment_2ea718ad-7d10-4222-a572-5d1cd61ecbe4). This difference may perhaps be explained by a difference in patterns of activity between these two bovids.
Both the grey rhebok and the common impala tend to ruminate in the open (as opposed to under cover of vegetation), where visual vigilance is enhanced during periods of lying down. However, the grey rhebok is cathemeral (https://www.oxfordreference.com/display/10.1093/oi/authority.20110803095555142 and https://treatment.plazi.org/GgServer/html/03F507139908FFB303D5F84BF585F2FE), whereas the common impala tends to lie down only at night (https://www.inaturalist.org/posts/49366-locomotory-and-postural-peculiarities-of-impalas-part-1#).
This suggests that the mutual monitoring of jaw-movements is performed usually in the dark in the common impala, versus often in the bright light of day in the grey rhebok. A pale feature is likely to be more conspicuous than a dark feature, by night.
What emerges from my scrutiny of the grey rhebok is two conspicuous features of colouration, as follows.
The caudal flag is large-scale, strikingly pale, activated by a combination of running and erection of the tail, and directed at potential predators, at distances of up to 0.5 km (https://www.inaturalist.org/observations/204059396 and https://www.inaturalist.org/observations/247276002).
The buccal semet is small-scale, subtlely dark, activated by a combination of cessation of chewing and synchronous swivelling of the ears, and directed intraspecifically, within the group, at distances of only up to 20 m (second photo in https://uk.inaturalist.org/observations/189700627).
Comentarios
https://www.inaturalist.org/journal/milewski/53696-a-succinct-photo-guide-to-subspecies-of-the-bush-duiker-sylvicapra-grimmia
https://www.inaturalist.org/posts/70156-subtle-diversification-of-buccal-semets-in-tragelaphin-antelopes-part-2#
A semet is a type of conspicuous pattern of colouration, in the same series as bleeze and flag.
A bleeze is a pattern so conspicuous that it makes the whole figure of the animal stand out, even at some distance. The adaptively important point is that this tends to preclude hiding from predators, even if the figure remains stationary.
A flag is a smaller-scale pattern, which becomes conspicuous at the scale of the whole figure only when moved. A typical location for flags is the tail.
A semet is also dependent on motion for its conspicuousness. However, it is so small-scale that it is only conspicuous at close range (less than 20 m for a large-bodied species, less than 10 m for a small-bodied species). Typical locations are ears and mouth.
Bleezes are mainly for long-range communication intraspecifically, the meaning being crude presence at a 'public' level.
Flags can communicate intraspecifically or with predators, the messages being social in some cases.
Semets communicate only intraspecifically, i.e. their messages are 'private', not 'public'.
Three photos to illustrate typical examples:
Bleeze https://naturerules1.fandom.com/wiki/Bontebok
Flag https://nuwejaars.com/did-you-know-the-grey-rhebok-is-creeping-up-the-endangered-list/
Semet https://www.gettyimages.ie/detail/video/an-eland-chews-its-cud-in-a-grassy-plain-in-south-stock-video-footage/1BT10342_0269
https://www.mindenpictures.com/stock-photo-grey-rhebok-pelea-capreolus-adult-close-up-of-front-hoof-undersurface-naturephotography-image80155895.html
For the buccal semet in Aepyceros melampus, see https://www.inaturalist.org/journal/milewski/89674-bleezes-flags-and-semets-in-the-adaptive-colouration-of-impalas-aepyceros-part-2#
Additional illustrations are:
https://www.gettyimages.com.au/detail/photo/two-male-impala-fighting-serengeti-national-park-royalty-free-image/75875136?phrase=african+impala&adppopup=true
https://www.gettyimages.com.au/detail/photo/female-impalas-on-the-alert-royalty-free-image/649107623?phrase=african+impala&adppopup=true
A fascinating read.
@bluehillescape
Many thanks for your appreciative words.
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