Also see https://www.inaturalist.org/journal/milewski/75748-adaptive-colouration-in-the-blesbok-damaliscus-pygargus-phillipsi-part-1-adults#
The bontebok (Damaliscus pygargus pygargus) is one of the most vividly-coloured wild ungulates on Earth.
However, its colouration has not previously been described coherently, relative to a conceptual framework based on adaptation. Furthermore, previous descriptions have been limited by the lack of suitable terms.
A reminder of the difference between a bleeze and a flag:
A bleeze is a large-scale feature of colouration, so bold that it makes the whole figure conspicuous even at a distance, and even when the animal is stationary.
A flag is a relatively small, or normally hidden, feature of colouration, which becomes conspicuous with motion.
The bontebok can be described as 'pied' (https://dictionary.cambridge.org/dictionary/english/pied). However, this is unsatisfactory, because
- it does not elucidate the functional organisation of the markings, and
- it does not distinguish between an adaptive design and that seen in pied individuals and breeds of domestic mammals (https://en.wikipedia.org/wiki/Piebald).
Various wild ungulates possess bleezes. However, what makes the bontebok unusual is that
In this account, I ignore the hues on the pelage of the bontebok, and consider only the tones (white, shades of grey, and black). This is because ungulates and their carnivoran predators do not perceive hues in the same ways as humans do.
OVERALL VIEWS
https://www.shutterstock.com/it/image-photo/walking-field-bontebok-mediumsized-generally-dark-484356043
https://www.inaturalist.org/observations/11251852
https://www.dreamstime.com/bontebok-damaliscus-pygargus-image129038646
https://www.agefotostock.com/age/en/details-photo/male-blesbok-baby-portrait-malolotje-swaziland-malolotje-nature-reserve-swaziland/ALM-BP5496/1
https://www.alamy.com/stock-photo-bontebok-damaliscus-pygargus-pygarus-de-hoop-nature-reserve-western-59178463.html?imageid=04F0A152-6CA3-4A65-9067-0364F1A5F09E&p=22059&pn=4&searchId=dca6a568865628209af3d56514ac1f36&searchtype=0
https://www.alamy.com/stock-photo-a-portrait-of-a-bontebok-antelope-in-south-africa-47742290.html?imageid=53CB260B-D3BB-4DD7-B43C-33C570D2CC36&p=158473&pn=1&searchId=abde0ec3b06a08943ffb523ff66f0db0&searchtype=0
https://www.alamy.com/bontebok-damaliscus-pygargus-in-game-park-in-south-africa-looking-over-its-shoulder-image245661564.html
https://www.dreamstime.com/bontebok-damaliscus-pygargus-image129038241
FACIAL BLEEZE
The conspicuously pale patch on the rostrum of adults of Damaliscus pygargus (https://www.inaturalist.org/taxa/42275-Damaliscus-pygargus) constitutes what is probably the clearest example of a facial bleeze, in any mammal.
In the bontebok, this patch is pure white, contesting with the dark ground-colour of the face and neck.
https://www.alamy.com/stock-photo-bontebok-blesbok-damaliscus-dorcas-standing-south-africa-47895393.html?imageid=A30EE114-7C91-437B-A4EE-F0E2062F42F1&p=33610&pn=6&searchId=ab384b9287a9a62b3e65ec702d7129d5&searchtype=0
https://www.alamy.com/stock-photo-bontebok-damaliscus-dorcas-at-bontebok-national-park-in-south-africa-38098213.html?imageid=2F8F0D32-30E5-4683-816A-5422AC0E0766&p=60598&pn=2&searchId=595b2aef109e6e3945fa2d85a0f44eb0&searchtype=0
https://www.alamy.com/stock-photo-bontebok-antelope-121112217.html?imageid=C030CB48-025D-450C-9009-14F6598A42A7&p=353301&pn=1&searchId=abde0ec3b06a08943ffb523ff66f0db0&searchtype=0
https://www.gettyimages.com.au/detail/photo/bontebok-damaliscus-dorcas-at-bontebok-national-royalty-free-image/523049311?phrase=bontebok&adppopup=true
https://www.alamy.com/stock-photo-a-pair-of-bontebok-antelopes-damaliscus-pygargus-dorcas-south-africa-85437994.html?imageid=8C3973B1-D533-4CA3-B604-463AAB155B2F&p=70019&pn=1&searchId=abde0ec3b06a08943ffb523ff66f0db0&searchtype=0
There is minimal variation in the facial bleeze, between the sexes, and among individuals. However, a few individuals show slight expansion at the base of the horns (https://www.alamy.com/stock-photo-blesbok-or-bontebok-damaliscus-dorcas-dorcas-syn-damaliscus-pygargus-78567725.html?imageid=967A4632-A630-4F22-84B5-7F50172963C0&p=183822&pn=2&searchId=dff47d9894fdcd82529cde91c8758091&searchtype=0).
Comparison with blesbok (https://www.inaturalist.org/journal/milewski/75748-adaptive-colouration-in-the-blesbok-damaliscus-pygargus-phillipsi-part-1-adults#):
The pale rostral patch on the face seems proportionately slightly smaller in the bontebok than in the blesbok, owing to what I perceive to be a difference in the size of the head.
Furthermore, in the bontebok the pale on the face tends not to expand with age, as it does in the blesbok (https://www.shutterstock.com/it/image-photo/blesbok-antelope-damaliscus-pygargus-standing-grass-1329980789 and https://www.inaturalist.org/journal/milewski/75871-a-previously-overlooked-expansion-of-the-bles-in-the-blesbok-signifying-full-maturity-beyond-adulthood#).
However, the facial bleeze is as well-developed in the bontebok as in the blesbok, because it seems exempt from the glandular staining seen in males of the blesbok (https://www.alamy.com/a-blesbok-antelope-damaliscus-pygargus-south-africa-image66899227.html?imageid=AA000BC4-7506-4457-96A0-B91FB81FCF46&p=70019&pn=1&searchId=a83f931787d9571cb0a25aca9f82d745&searchtype=0 and https://www.alamy.com/stock-photo-portrait-of-a-blesbok-antelope-damaliscus-pygargus-south-africa-59076858.html?imageid=9FB1F37F-B684-4FE4-A0A4-9A9AC4C55B63&p=70019&pn=1&searchId=a83f931787d9571cb0a25aca9f82d745&searchtype=0).
LATERAL BLEEZE
The lateral bleeze of the bontebok is, proportionately, one of the largest bleezes seen in any ungulate. It extends from the withers and the back to the ventral edge of the thorax and the belly. It also Includes the posterior surface of the upper foreleg, and the anterior surface of the hindleg just below the knee (https://www.inaturalist.org/observations/3985830 and https://www.inaturalist.org/observations/38720040).
The pattern is one of horizontal banding on a broad design, in which the upper and lower panels are pale, with the flanks forming a dark median panel.
However, an important qualification is that the upper panel appears pale partly owing to sheen. Its conspicuousness thus depends on illumination.
https://www.dreamstime.com/stock-photo-bontebok-field-looking-perfect-medium-sized-generally-dark-brown-antelope-prominent-wide-white-image77512668
https://www.agefotostock.com/age/en/details-photo/bontebok-damaliscus-dorcas-dorcas-de-hoop-reserve-south-africa-africa/RHA-743-243/1
https://www.agefotostock.com/age/en/details-photo/bontebok-damaliscus-dorcas-dorcas-bovidae-antelope-mammal-animal-cape-of-good-hope-national-park-south-africa-africa/H44-10978980/1
https://www.alamy.com/stock-photo-bontebok-damaliscus-pygargus-pygarus-at-the-base-of-the-langeberg-59175692.html?imageid=FE1F6904-9478-4A53-8A24-5E8D481C7E90&p=22059&pn=3&searchId=6d8a7f95c3b92935b5e8c1d5d07d00ff&searchtype=0
https://www.alamy.com/bontebok-in-the-bontebok-national-park-swellendam-western-cape-south-africa-image342638387.html?imageid=5C55BBE5-4514-4B34-AEDA-ACC05175FD68&p=22337&pn=3&searchId=6d8a7f95c3b92935b5e8c1d5d07d00ff&searchtype=0
https://www.inaturalist.org/observations/113481398
https://www.inaturalist.org/observations/115575412
https://www.inaturalist.org/observations/80586175
https://www.alamy.com/stock-photo-bontebok-antelope-114770771.html?imageid=0A5EB348-1301-4DC3-B4AD-F99E5785ACEA&p=353301&pn=4&searchId=29c543d132619cf1217ce0a23a3945c6&searchtype=0
https://www.gettyimages.com.au/detail/photo/bonteboks-grazing-royalty-free-image/541049632?phrase=blesbok&adppopup=true
The design of the lateral bleeze of the bontebok is such that in bright sunlight at midday, when the ventral white is inconspicuous owing to shading, the dorsal sheen is 'switched on' owing to sheen (https://www.inaturalist.org/observations/27337501 and https://www.inaturalist.org/observations/27289820).
Comparison with blesbok:
A categorical difference is that, unlike the bontebok, the blesbok lacks a lateral bleeze.
The pelage of the flanks is not necessarily any paler in the blesbok than in the bontebok. However, the differences are that, in the blesbok,
- the dorsal sheen is poorly-developed,
- the pale ventral pelage is restricted, and reduced to a combination of countershading and an abdominal flag, and
- the white anterior edge of the hindleg, just below the knee, is narrower than in the bontebok.
Furthermore,
- the abdominal flag of the blesbok (https://www.inaturalist.org/observations/130975314) is subsumed within a broader white, ventral panel, so that the bontebok lacks an abdominal flag, and
- the ulnar dark/pale contrast of the blesbok - although just as clear in the bontebok - becomes an anterior extension of the ventral white of the thorax, so that the bontebok lacks an ulnar flag.
PYGAL BLEEZE
The pygal bleeze of the bontebok is about as large as the facial bleeze, but located at the opposite pole of the figure (https://www.alamy.com/stock-photo-a-pair-of-bontebok-antelopes-damaliscus-pygargus-dorcas-south-africa-85437994.html?imageid=8C3973B1-D533-4CA3-B604-463AAB155B2F&p=70019&pn=1&searchId=abde0ec3b06a08943ffb523ff66f0db0&searchtype=0 and https://www.dreamstime.com/stock-photo-bontebok-antelopes-pair-damaliscus-pygargus-dorcas-south-africa-image56821286).
The pale posterior feature of the bontebok
- extends from the rump to the tail-stalk and the buttocks immediately adjacent to the anus and perineum,
- is largely white, and
- contrasts with the dark pelage around it.
https://www.inaturalist.org/observations/80650367
https://www.inaturalist.org/observations/57714646
https://www.alamy.com/stock-photo-a-bontebok-damaliscus-pygargus-in-the-bontebok-national-park-near-173731691.html?imageid=75A1E59F-9E57-44B2-8852-46B50B30641E&p=77702&pn=6&searchId=d7a105eea18d32dc6155ddf3de390174&searchtype=0
https://www.inaturalist.org/observations/53977868
Comparison with blesbok:
The pygal flag of the blesbok bears the same relationship to the pygal bleeze of the bontebok as the abdominal flag of the blesbok (https://www.alamy.com/blesbok-in-savannah-south-africa-image151809725.html?imageid=3B900166-AC81-4937-96B6-1CA462655F62&p=703420&pn=10&searchId=26e0b007d6f642f3bb26a9cc6900a5cf&searchtype=0) bears to the white ventral panel of the lateral bleeze of the bontebok.
Furthermore, the pygal flag of the blesbok depends mainly on sheen effects, whereas the pygal bleeze of the bontebok depends mainly on the depigmentation of the pelage.
PEDAL FLAG
The lower legs of adults of the bontebok are largely white, making them conspicuous in locomotion.
https://www.inaturalist.org/observations/127885416
https://www.dreamstime.com/stock-photo-bontebok-antelope-running-velt-image65832109
https://www.alamy.com/bontebok-are-the-colourful-bok-of-south-africa-image228097057.html?imageid=7AAA4F55-17D1-423B-BF91-57470077439F&p=641396&pn=1&searchId=25869b8bb63848eab5bc67f94e53a679&searchtype=0
https://www.alamy.com/stock-photo-bontebok-antelope-17066360.html?imageid=AFA96455-2E25-41C8-91B0-1B7B94A1EA3C&p=4877&pn=1&searchId=abde0ec3b06a08943ffb523ff66f0db0&searchtype=0
https://www.alamy.com/stock-photo-bontebok-antelope-17066531.html?imageid=87C5DBB8-BEBE-41B5-A483-7EF7427CDD10&p=4877&pn=3&searchId=8e369419b3a055a70321251ed7501637&searchtype=0
Comparison with blesbok:
Both the bontebok and the blesbok possess a pedal flag. However, this feature is better-developed in the bontebok, in which
- the white pelage is more extensive, particularly on the outer surfaces of the lower legs, and
- there is less individual variation.
LACK OF AURICULAR FLAG
In the bontebok, there is a considerable sheen on the short, sparse pelage on the posterior surface of the ear pinnae. In some.views this produces a noticeably pale aspect to the back-of-ear.
However, after perusing hundreds of photos, I have not found this effect to be strong or consistent enough to qualify as an auricular flag. It is better-regarded as an incipient/residual version of the pattern in the blesbok (see below).
https://www.agefotostock.com/age/en/details-photo/bontebok-antelope/ESY-001254401/1
https://www.alamy.com/bontebok-antelope-damaliscus-pygargus-dorcas-south-africa-image156518411.html?imageid=FCBE4CA1-DBF7-4706-9D07-20EC04F83E25&p=476215&pn=1&searchId=abde0ec3b06a08943ffb523ff66f0db0&searchtype=0
https://www.inaturalist.org/observations/84604329#data_quality_assessment
https://www.alamy.com/stock-photo-bontebok-antelope-bontebok-national-park-swellendam-overberg-western-139221333.html?imageid=C6DAAADA-77BC-4140-AC0F-6ACCCDFABC9D&p=10587&pn=1&searchId=abde0ec3b06a08943ffb523ff66f0db0&searchtype=0
https://www.alamy.com/stock-photo-at-the-peaceful-and-quiet-bontebok-south-african-park-two-antelopes-32166159.html?imageid=143BD930-B2D0-433C-A84D-753F5F9CE194&p=94763&pn=1&searchId=abde0ec3b06a08943ffb523ff66f0db0&searchtype=0
https://www.inaturalist.org/observations/115550547
https://www.alamy.com/bontebok-walking-image647406.html?imageid=FF50FFAE-284D-4D2A-B72E-56FEB151968F&p=4877&pn=4&searchId=29c543d132619cf1217ce0a23a3945c6&searchtype=0
Comparison with blesbok:
The ear pinnae seem identical in size and pigmentation in the bontebok and the blesbok.
However, perusal of hundreds of photos shows that the sheeny quality of the posterior surface is better-developed in the blesbok (https://www.agefotostock.com/age/en/details-photo/blesboks-damaliscus-pygargus-phillipsi-two-adults-standing-in-open-grassland-alert-mountain-zebra-national-park-eastern-cape-south-africa-africa/ZI6-3207754/1 and https://www.agefotostock.com/age/en/details-photo/bontebok-south-africa/FHR-88889-04144-075/1) than in the bontebok.
This, together with the relatively poorly-developed sheen on the withers and back, qualifies the blesbok, in my view, for an auricular flag (https://www.inaturalist.org/journal/milewski/75905-unlike-the-blesbok-the-bontebok-damaliscus-pygargus-pygargus-lacks-an-auricular-flag#).
The lack of an auricular flag in the bontebok is significant evolutionarily. This is because all of the postcranial pale features are better-developed in the bontebok than in the blesbok.
LACK OF CAUDAL FLAG
The tail of the bontebok is large enough to be conspicuous, particularly because the white tail-stalk contrasts with the black tail-tassel (https://www.agefotostock.com/age/en/details-photo/bontebok-antelope/ESY-008277294/1). However, in terms of colouration, the tail is subsidiary to the pygal bleeze, which is the more conspicuous feature.
Furthermore, the bontebok displays its tail only in a few social contexts (e,g. courtship, https://www.alamy.com/stock-photo-bontebok-blesbok-antelope-damaliscus-damaliscus-dorcas-bovidae-male-36820913.html?imageid=D177A04E-5BB2-42A8-8538-B839CCC540E6&p=26096&pn=5&searchId=78e3c36a9917c21e07efdd82f4c4fbc0&searchtype=0), and not in alarm or fleeing. It does not seem to wag/swish the tail except in reaction to insects.
As a result, the bontebok does not, in my view, qualify for a caudal flag.
https://www.alamy.com/stock-photo-africa-south-africa-west-coast-national-park-bontebok-antelope-damaliscus-56090059.html?imageid=7A7DA0E1-EDD1-47FE-9168-012DC072ABCE&p=183410&pn=1&searchId=abde0ec3b06a08943ffb523ff66f0db0&searchtype=0
Comparison with blesbok:
The tail of the blesbok differs from that of the bontebok in that
However, the blesbok likewise fails to qualify for a caudal flag. This is because
- the dark/pale contrast on the tail is less, not more, developed than in the bontebok, and
- the blesbok, like the bontebok, 'downplays' the tail in terms of demonstrations, particularly in the context of anti-predator behaviour.
DISCUSSION
The colouration of adults of the bontebok is configured in such a way that the figure is conspicuous, regardless of whether
to be continued in https://www.inaturalist.org/journal/milewski/76336-adaptive-colouration-in-the-bontebok-damaliscus-pygargus-pygargus-part-2-infants-juveniles-and-adolescents#...