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INTRODUCTION
The roan antelope (Hippotragus equinus, https://www.inaturalist.org/taxa/42332-Hippotragus-equinus) is an unusually widespread species of African bovid.
It spans the continent from far-West Africa to Eritrea, and southward to subtropical South Africa (https://upload.wikimedia.org/wikipedia/commons/e/e1/Roan_Antelope_Hippotragus_equinus_distribution_map.png and https://www.researchgate.net/figure/21-The-geographic-range-of-roan-antelope-in-the-1800s-Du-Plessis-1969-and-today-IUCN_fig2_278698501).
Given this vast range, we might expect that distinct subspecies occur (https://www.researchgate.net/figure/The-distribution-of-roan-antelope-in-Africa-Anonymous-2006-with-subspecies-delineations_fig1_290478320).
However, Groves and Grubb (2011, pages 198-201, including Table 53, in https://zmmu.msu.ru/files/%D0%91%D0%B8%D0%B1%D0%BB%D0%B8%D0%BE%D1%82%D0%B5%D0%BA%D0%B0%20%D0%9F%D0%B0%D0%B2%D0%BB%D0%B8%D0%BD%D0%BE%D0%B2%D0%B0/grubb-groves-2011_taxonomy_ungulates.pdf), in their taxonomic revision of all the ungulates of the world, failed to verify any subspecies of the roan antelope.
Their revision was based on museum specimens, ignoring photographic evidence such as that now available electonically.
Factors that have retarded/complicated verification of subspecies of the roan antelope in the past include:
MY APPROACH: HIPPOTRAGUS EQUINUS KOBA Vs HIPPOTRAGUS EQUINUS EQUINUS/COTTONI
My perusal of observations in iNaturalist, and photos elsewhere on the Web, has shown certain fairly consistent differences, particularly in colouration, between the subspecies at the extremes of the geographical range.
Most of these subspecific differences were unbeknownst to Groves and Grubb (2011), and to this day no other taxonomist seems to have noticed them.
Please bear in mind that all of the following are subject to individual variation - which is more important than sexual differences.
The ground-colour of the West African subspecies (H. e. koba) tends to be relatively bright-hued (Groves and Grubb 2011), with minimal grizzling and relatively short pelage. The hues are difficult to compare photographically, owing to vagaries of illumination and technique.
However, the extreme length and pallor of the beard, shown in the following captive specimens of Hippotragus equinus equinus/cottoni, are - as far as I know - never seen in H. e. koba:
adult female https://www.alamy.com/a-close-up-profile-portrait-of-a-roan-antelope-hippotragus-equinus-it-shows-the-head-and-neck-together-with-large-antlers-image429376262.html?imageid=51F04DDC-EFAB-4CE4-B304-781CE27CA2CB&p=224192&pn=3&searchId=907fcf51e051a28f272ac53440026b09&searchtype=0
juvenile https://www.shutterstock.com/image-photo/close-roan-antelope-beautiful-herbivore-1713171139.
Another fairly obvious difference is that, in the West African subspecies (H. e. koba) relative to the southern African subspecies (H. e. equinus/cottoni), dark pelage can be
Let us now examine, in particular, the tendency - consistently visible in photos - for the dark pelage on the face to be more extensive in the West African than in the southern African subspecies.
HOW TO SCRUTINISE THE FACE
A conspicuously pale feature to focus on is the preorbital tuft, located anterior and ventral to the eye.
Dear Reader, can you spot the difference between https://www.inaturalist.org/observations/61091684 and https://www.shutterstock.com/image-photo/roan-antelope-hippotragus-equinus-mammal-1521109436, in the size and shape of the preorbital tuft?
In infancy in both subspecies, the nucleus of the dark 'mask' appears, on
From this nucleus the darkness spreads (https://www.alamy.com/roan-antelope-hippotragus-equinus-female-both-sexes-have-long-curved-scimitar-shaped-ribbed-horns-black-and-white-facial-markings-long-ears-image398724544.html?imageid=9C93FB19-3D12-4AE0-BF38-AAFEA8A9CAEE&p=77483&pn=1&searchId=a5e37347722537f38c2dc3ebc9b10b9f&searchtype=0), according to age and sex,
This spread tends ultimately to be greater in H. e. koba than in H. e. equinus/cottoni; in fully mature males of the former, the face becomes mainly black.
In H. e. koba, even the following are subject to discoloration in some individuals:
VARIOUS DETAILED DIFFERENCES
Now, let us approach various facial and other features in detail:
In the West African subspecies,
PAIRWISE PHOTO-COMPARISONS
In the following pairwise comparisons, I illustrate the above aspects of ssp. koba of West Africa relative ssp. equinus/cottoni of southern Africa:
dark/pale pattern on the face
https://www.inaturalist.org/observations/73846242
https://en.wikipedia.org/wiki/Portal:Africa/Featured_picture#/media/File:Roan_antelope_(Hippotragus_equinus_koba).jpg
https://www.gettyimages.ae/detail/news-photo/hippotragus-equinus-roan-antelope-news-photo/1198611040?adppopup=true
https://www.naturepl.com/stock-photo-roan-antelope-hippotragus-equinus-male-in-profile-fathala-reserve-nature-image01387428.html
https://www.inaturalist.org/observations/151288128
https://www.alamy.com/stock-photo-roan-antelope-hippotragus-equinus-rserve-de-bandia-senegal-31330979.html?imageid=D9641CA0-1D7F-47C7-B77A-9C42739A2A9E&p=22059&pn=2&searchId=414e17011db666d97683dcc728f39b91&searchtype=0
https://www.inaturalist.org/observations/181196418
https://www.inaturalist.org/observations/149568372
VERSUS
https://www.alamy.com/stock-photo-roan-antelope-hippotragus-equinus-in-profile-close-up-84710327.html
https://www.alamy.com/roan-antelope-hippotragus-equinus-portrait-south-africa-image255403284.html?imageid=558E7226-6400-4BC9-9CAA-EC91E04DFF0F&p=851122&pn=1&searchId=d3350f0b953b17636829cec591564ee8&searchtype=0
https://www.inaturalist.org/observations/8255165
https://www.inaturalist.org/observations/8221501
https://www.inaturalist.org/observations/173923530
https://www.inaturalist.org/observations/105211286
https://www.inaturalist.org/observations/12434566
https://howieswildlifeimages.files.wordpress.com/2022/06/2021-09-18_m1x-p9183992_dxo.jpg
https://www.alamy.com/portrait-of-a-male-roan-antelope-hippotragus-equinussouth-africa-image68948756.html?imageid=E726C045-9A48-4FBD-B104-A87D09D7790C&p=3864&pn=1&searchId=d3350f0b953b17636829cec591564ee8&searchtype=0
pattern on flanks and belly
https://www.inaturalist.org/observations/5280585
https://www.inaturalist.org/observations/103681190
https://www.inaturalist.org/observations/156041605
VERSUS
https://www.inaturalist.org/observations/12434566
https://www.inaturalist.org/observations/5635964
pattern on hindquarters, from rump to buttocks and tail
https://www.inaturalist.org/observations/9979094
https://www.inaturalist.org/observations/58642342
https://www.inaturalist.org/observations/66772812
https://www.inaturalist.org/observations/109062982
https://www.inaturalist.org/observations/25801218
VERSUS
https://www.inaturalist.org/observations/188061458
https://www.istockphoto.com/photo/a-rare-roan-antelope-in-natural-habitat-south-africa-gm1628183210-532254021?phrase=roan+antelope
https://www.shutterstock.com/image-photo/stunning-roan-antelope-hwange-national-park-1186020874
https://www.inaturalist.org/observations/36915943
dark/pale pattern on feet
https://www.inaturalist.org/observations/6793183
https://www.inaturalist.org/observations/91165756
https://www.inaturalist.org/observations/103681181
https://www.inaturalist.org/observations/195546644
https://www.inaturalist.org/observations/103681226
VERSUS
https://www.inaturalist.org/observations/138018015
https://howieswildlifeimages.files.wordpress.com/2022/05/2021-09-18_m1x-p9183957_dxo.jpg
DISCUSSION
One of the most remarkable facts about the roan antelope is how little it varies across a vast - and somewhat disjunct - geographical distribution.
As a result, populations more than ten thousand kilometres apart (Senegal, https://upload.wikimedia.org/wikipedia/commons/3/3c/Senegal_%28orthographic_projection%29.svg, versus North West Province of South Africa, https://upload.wikimedia.org/wikipedia/commons/f/fc/North_West_in_South_Africa.svg) appear so similar, in photos, that their subspecific distinction could be doubted.
However, there are two differences that, although slight, may as well be categorical in the overall impression they leave on the human viewer. These are
Were it the historical case that the species had become well-known in West Africa (francophone) before South Africa (anglophone), it would probably not have been given the name 'roan antelope' (French: 'antilope rouanne'). A more apt name for H. e. koba might have been 'antilope masquee' ('masked antelope').
This is because the ground-colour in West Africa is not reminiscent of roan colouration in the domestic horse (https://upload.wikimedia.org/wikipedia/commons/f/fc/Red_roan_Quarter_Horse.jpg and https://en.wikipedia.org/wiki/Roan_(color) and https://www.deviantart.com/hunter-raider/art/Coat-Color-Guide-2023-Roan-965502025).
Perhaps the first diagnostic feature to focus on, in most photos, is the preorbital tuft, which is whitish, surrounded by dark, short pelage.
The following, of H. e. equinus/cottoni, show the maximum development of the whitish preorbital tuft:
https://www.shutterstock.com/image-photo/closeup-resting-roan-antelope-hippotragus-equinus-2177832261 and https://www.chesterzoo.org/our-zoo/animals/roan-antelope/.
The following, of H. e. koba, shows how much smaller this feature is in the West African subspecies:
https://www.shutterstock.com/image-photo/roan-antelope-hippotragus-equinus-savanna-found-1391679644.
The difference in the size of the preorbital tuft is
However, it is noteworthy that
The West African subspecies is larger-bodied than the southern African subspecies. Mature males weigh respectively about 300 kg and 250 kg (https://sportsafield.com/2021/its-a-roan/).
To the trained eye, this difference is hinted at by the proportionately slightly smaller post-withers mane-tuft in H. e. koba (https://www.inaturalist.org/journal/milewski/87946-has-the-post-withers-mane-tuft-of-hippotragus-evolved-in-mimicry-of-oxpeckers-buphagus-part-1#).
The caudo-ischial flag (https://www.inaturalist.org/posts/85734-oryx-in-posteriolateral-view-an-interspecies-comparison-of-adaptive-colouration#) is somewhat more poorly-developed in H. e. koba than in H. e. equinus/cottoni.
The converse is true in the case of an incipient/residual pedal flag. This pattern is slight, even in H. e. koba.
Comentarios
What is the dark feature on the lower lip?
https://www.inaturalist.org/observations/103681241
Sheen on the neck of the southern African subspecies:
https://www.inaturalist.org/observations/67879483
https://www.inaturalist.org/observations/57812246
The following individual shows the maximum development of dark/pale contrast on the hindquarters in posteriolateral view, in the West African subspecies:
https://www.inaturalist.org/observations/25801218
Does the mane begin farther from the occiput than in the southern African subspecies?
https://www.inaturalist.org/observations/17942082
The following show the typical appearance of adult males in the West African subspecies:
https://www.onlinehuntingauctions.com/CLUB-FAUNE-7-Day-West-African-Savannah-Buffalo-OR-Roan-Antelope-Hunt-for-One-Hunter-and-One-Non-Hun_i43579639
https://www.africaimagelibrary.com/-/galleries/west-africa/senegal/-/medias/c9e38cc0-2120-11e0-96ba-bd32c8fafe4a-roan-antelope-hippotragus-equinus-reserve-de-bandia-senegal
https://www.inaturalist.org/guide_taxa/841811
https://www.africaimagelibrary.com/-/galleries/west-africa/senegal/-/medias/256348d8-2121-11e0-beca-75e7a1c29f27-roan-antelope-hippotragus-equinus-reserve-de-bandia-senegal
The following video contains some clear depictions:
https://www.youtube.com/watch?app=desktop&v=Cgy0R938mdU
https://www.sa-venues.com/wildlife/wildlife_roan_antelope.php
The following shows flehmen in mature male of the West African subspecies. Please note the complete loss of the dark feature at the apex of the ear pinna:
https://www.facebook.com/photo/?fbid=705715644918042&set=a.553610316795243&locale=es_ES
The following shows the typical appearance of adolescent/adult females in the southern African subspecies:
https://www.alamy.com/roan-antelope-hippotragus-equinussavanna-antelope-found-in-west-central-east-and-southern-africa-detail-portrait-of-antelope-head-with-big-ears-image466880633.html
https://www.gettyimages.ae/detail/photo/close-up-roan-antelope-royalty-free-image/1320270979?phrase=these+roan+antelope&adppopup=true
The following is unusual in showing clearly an infant individual (probably about one month old) of the West African subspecies:
Please note a) that dark on the tail is more precocial than that on the face, and b) the pale preorbital tuft is small.
https://www.africaimagelibrary.com/media/38bfbf0c-2120-11e0-8287-8bc0e7630dc0-roan-antelope-hippotragus-equinus-reserve-de-bandia-senegal
EXTREMELY CLEAR DEPICTION OF COLOURATION
https://www.reserveafricainesigean.fr/en/animals/roan-antelope/
The following, in Kruger National Park, is an exceptionally clear illustration of 'roan colouration', similar to that in the domestic horse:
https://howieswildlifeimages.files.wordpress.com/2022/05/2021-09-18_m1x-p9183964_dxo.jpg
This plus subsequent comments:
Adolescent males in Kruger National Park:
https://howieswildlifeimages.files.wordpress.com/2022/05/2021-09-18_m1x-p9183968_dxo.jpg
https://howieswildlifeimages.files.wordpress.com/2022/05/2021-09-18_m1x-p9183985_dxo.jpg
https://howieswildlifeimages.files.wordpress.com/2022/05/2021-09-18_m1x-p9188214.jpg
https://howieswildlifeimages.files.wordpress.com/2022/05/2021-09-18_m1x-p9188223_dxo.jpg
Clear depiction of facial profile:
https://www.whiteoakwildlife.org/wildlife/roan-antelope/
https://www.flickr.com/photos/annette_rumbelow/49918940798
https://www.gettyimages.ae/detail/photo/portrait-of-a-male-roan-antelope-south-africa-royalty-free-image/541054642?phrase=these+roan+antelope&adppopup=true
https://www.gettyimages.ae/detail/photo/portrait-of-a-male-roan-antelope-south-africa-royalty-free-image/541054616?phrase=these+roan+antelope&adppopup=true
Six week-old infants of the southern African subspecies:
https://www.gettyimages.ae/detail/photo/roan-antelope-young-calves-south-africa-royalty-free-image/541054438
https://www.gettyimages.ae/detail/photo/roan-antelope-female-and-calves-south-africa-royalty-free-image/541054578?phrase=these+roan+antelope&adppopup=true
A senile male individual of the southern African subspecies:
https://www.gettyimages.ae/detail/photo/roan-antelope-royalty-free-image/622152390?phrase=these+roan+antelope&adppopup=true
Malawi:
https://www.gettyimages.ae/detail/news-photo/roan-antelope-in-the-grasslands-of-the-nyika-plateau-nyika-news-photo/589466518?adppopup=true
https://www.gettyimages.ae/detail/news-photo/roan-antelope-in-the-grasslands-of-the-nyika-plateau-nyika-news-photo/589466508?adppopup=true
https://www.gettyimages.ae/detail/news-photo/roan-antelope-in-the-grasslands-of-the-nyika-plateau-nyika-news-photo/589313878?adppopup=true
Adult male of southern African subspecies:
https://www.gettyimages.ae/detail/photo/portrait-of-a-roan-antelope-south-africa-royalty-free-image/1043283214?phrase=these%2Broan%2Bantelope
Adult male of ?West African subspecies:
https://www.gettyimages.ae/detail/photo/roan-antelope-royalty-free-image/579764051?phrase=these+roan+antelope&adppopup=true
https://www.gettyimages.ae/detail/photo/roan-antelope-royalty-free-image/1145467580?phrase=these+roan+antelope&adppopup=true
https://www.gettyimages.ae/detail/photo/roan-zebra-savuti-chobe-botswana-royalty-free-image/1149416207?phrase=these+roan+antelope&adppopup=true
https://www.gettyimages.ae/detail/photo/roan-antelope-royalty-free-image/980035828?phrase=these+roan+antelope&adppopup=true
The following nicely illustrates how small the whitish preorbital tuft is in the West African subspecies:
https://www.africaimagelibrary.com/media/626ef2be-2120-11e0-a7cb-6ff07bbaab0d-roan-antelope-hippotragus-equinus-reserve-de-bandia-senegal
https://sportsafield.com/2021/its-a-roan/
https://www.shutterstock.com/image-photo/endangered-roan-antelope-south-african-bush-2168068715
Infant just a few weeks old, showing e.g. the precociality of the post-withers mane-tuft:
https://www.shutterstock.com/image-photo/roan-antelope-hippotragus-equinus-baby-188068073
https://www.shutterstock.com/image-photo/roan-antelope-hippotragus-equinus-baby-188068052
https://animaldiversity.org/accounts/Hippotragus_equinus/pictures/collections/contributors/eva_hejda/pferdeantilope/
https://en.wikipedia.org/wiki/Preorbital_gland
https://eol.org/pages/331075/media
KOBUS DEFASSA
https://www.alamy.com/the-head-and-face-of-a-wild-male-waterbuck-pictured-in-the-kidepo-valley-national-park-northern-uganda-image506105245.html?imageid=1EF19C74-372F-43DC-A268-57491E403E74&p=390716&pn=1&searchId=a5e37347722537f38c2dc3ebc9b10b9f&searchtype=0
https://www.alamy.com/the-head-and-face-of-a-wild-male-waterbuck-pictured-in-the-kidepo-valley-national-park-northern-uganda-image506105912.html?imageid=5957F78D-2D0E-4FA6-A7B0-3DCD2709C0FC&p=390716&pn=1&searchId=a5e37347722537f38c2dc3ebc9b10b9f&searchtype=0
The following show the range of variation, among ruminants, in orbital width:
https://www.alamy.com/stock-photo-face-portrait-antelope-114743452.html?imageid=05570D68-D012-4839-B3D8-EDC02EDFBA64&p=335153&pn=1&searchId=a5e37347722537f38c2dc3ebc9b10b9f&searchtype=0
https://www.alamy.com/wildlife-portrait-of-a-prong-horned-antelope-in-south-dakota-united-image62287810.html?imageid=7E9C0BC9-163D-4B3E-B00E-483BD323DA1B&p=178246&pn=1&searchId=a5e37347722537f38c2dc3ebc9b10b9f&searchtype=0
https://www.inaturalist.org/observations/97540700
Here are unusual photos of infants of the roan antelope.
In the following (?4 weeks old), please note the suggestion of a fibular/tibial flag, and the fact that the dark apical ear-tufts have yet to appear:
https://www.flickr.com/photos/gilli8888/34599741036?
In the following (?6 weeks old), please note the precociality of the darknes on the tail:
https://www.veronasera.it/cronaca/parco-natura-viva-top-five-nascite-particolari-27-settembre-2016.html
Dark skin on neck
https://www.inaturalist.org/observations/2872343
https://www.google.com/search?q=roan+antelopes+Chester+zoo&sca_esv=598330860&tbm=isch&sxsrf=ACQVn08kIChSMHqCXT6NJmBH-6RR66Buaw:1705221354028&source=lnms&sa=X&ved=2ahUKEwjQxMfIvNyDAxXMcGwGHWkUAYUQ0pQJegQIBxAE&biw=904&bih=494&dpr=3#imgrc=NoT0LGLqeVvYGM&vwlns=WyIwQ0JRUWg2Y0dhaGNLRXdqZ3M2LWR2ZHlEQXhVQUFBQUFIUUFBQUFBUUpRIl0=&lns=W251bGwsbnVsbCxudWxsLG51bGwsbnVsbCxudWxsLG51bGwsIkVrY0tKRGs1WVRJd01EUTNMVFptTjJZdE5HWXhNaTA0T1dVNExUQTBOVGxqWVRReU9XWmtOaElmTURaSWJuQTBja0ZPZFZsVGMwWldkMUYzZG5veE4wVXlaRlpXTWpCQ1p3PT0iLG51bGwsbnVsbCxudWxsLDEsbnVsbCxbbnVsbCxudWxsLFswLDAsMTAwMDAwLDEwMDAwMF1dXQ==
An unusually clear illustration of the length of the tail at various ages in Loxodonta africana, which has a shorter tail than in Elephas maximus:
https://www.everypixel.com/q/animal-breeding?image_id=10845680870747623680
https://www.researchgate.net/publication/316165442_Focus_on_the_Roan_AntelopeHippotragus_equinus
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